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ArticleTitle The Carnian/Norian boundary succession at Berlin-Ichthyosaur State Park (Upper Triassic, central NV, USA) Article Sub-Title Article CopyRight Paläontologische Gesellschaft (This will be the copyright line in the final PDF) Journal Name Paläontologische Zeitschrift Corresponding Author Family Name Balini Particle Given Name Marco Suffix Division Dipartimento di Scienze della Terra “Ardito Desio” Organization University of Milan Address Via Mangiagalli 34, Milan, 20133, Italy Email [email protected]

Author Family Name Jenks Particle Given Name James F. Suffix Division Organization Address 1134 Johnson Ridge Lane, West Jordan, UT, 84084, USA Email Author Family Name Martin Particle Given Name Riccardo Suffix Division Dipartimento di Scienze della Terra “Ardito Desio” Organization University of Milan Address Via Mangiagalli 34, Milan, 20133, Italy Email Author Family Name McRoberts Particle Given Name Christopher A. Suffix Division Geology Department Organization State University of New York at Cortland Address P.O. Box 2000, Cortland, 13045, USA Email Author Family Name Orchard Particle Given Name Michael J. Suffix Division Organization Natural Resources Canada-Geological Survey of Canada Address 1500-605 Robson St., Vancouver, BC, V6B 5J3, Canada Email Author Family Name Silberling Particle Given Name Norman J. Suffix Division Organization US Geological Survey Address Menlo Park, USA Email

Received 3 December 2013 Schedule Revised Accepted 22 August 2014 Abstract The Upper Carnian-Lower Norian (Upper Triassic) Luning Formation at Berlin-Ichthyosaur State Park (BISP) in central NV (USA) has been sampled using for the first time the bed-by-bed approach for ammonoids, pelagic bivalves, and conodonts, more than 60 years after its first description by Silberling (U.S. Geological Survey Professional Paper 322: 1–63, 1959). BISP is historically important for the definition of the uppermost Carnian of the North American Triassic chronostratigraphic scale and is known worldwide as one of the most important ichthyosaur fossil-Lagerstätte because of its extraordinary record of 37 articulated, large-sized specimens of Shonisaurus popularis. Nearly 190 ammonoids were collected from two stratigraphic sections, documenting all the latest Carnian to Early Norian ammonoid faunas previously described by Silberling. Halobiids were collected from five levels, and the first report of conodonts from BISP includes faunas from 13 levels. The ~340-m thick Brick Pile section, the most complete in the study area, includes the uppermost Carnian Macrolobatus zone, which provides conodont faunas of the lower primitia zone and Halobia septentrionalis. The 200-m thick lowermost Norian Kerri zone, which begins 52 m above the Macrolobatus zone, yields conodonts of the upper primitia zone in its lower part, together with H. cf. beyrichi and H. cf. selwyni. The ichthyosaur-bearing interval, whose stratigraphic position has been interpreted quite differently by previous authors, is documented in the uppermost Carnian Macrolobatus zone and is characterized by rich Tropites-dominated ammonoid faunas and by the onset of Halobia. All models proposed by various workers to explain the unusual ichthyosaur record are discussed and an additional explanation for the main ichthyosaur- bearing bed is proposed. The new hypothesis is that a harmful algal bloom (HAB) may have been the trigger for the mass mortality recorded in this level. Although the C/N boundary in the Brick Pile section lies within a 52 m interval that presently lacks paleontologic data, this succession is included in a small group of sections that are expected to contribute to the definition of the GSSP of the Norian stage. Correlation of the Brick Pile with the best Carnian/Norian sections in northeastern British Columbia is discussed. Compared to the British Columbia Juvavites cove and the GSSP candidate Black Bear Ridge sections, the Brick Pile section exhibits an ammonoid and Halobia record that is slightly more similar to that of the Tethyan sections. Correlation of the Brick Pile with the second GSSP candidate Pizzo Mondello (Sicily, Italy) well demonstrates the significant problems encountered in calibration of the Tethyan and North American scales. Kurzfassung Die oberkarnische bis unternorische (Späte Trias) Luning Formation im Berlin-Ichthyosaur State Park (BISP) in Zentral-Nevada (USA) wurde mehr als 60 Jahre nach der ersten Beschreibung durch Silberling (U. S. Geological Survey Professional Paper 322: 1–63, 1959) erstmals bankweise auf Ammoniten, pelagische Bivalven und Conodonten untersucht. Der BISP ist von historischer Bedeutung für die Definition des obersten Karn in der chronostratigraphischen Tabelle der nordamerikanischen Trias und ist eine der weltweit bekanntesten Ichthyosaurier Fossillagerstätten, ausgezeichnet durch den außergewöhnlichen Fund von 37 artikulierten, großen Exemplaren von Shonisaurus popularis. Nahezu 190 Ammoniten aus zwei stratigraphischen Abschnitten belegen oberstes Karn bis Unternor, wie schon früher durch Silberling beschrieben. Halobiiden liegen aus fünf Niveaus vor, und der Erstnachweis von Conodonten im BISP beinhaltet Faunen aus 13 Niveaus. Der ca. 340 m mächtige Brick Pile Aufschluss ist der kompletteste im untersuchten Gebiet. Er schließt die rund 20 m mächtige Macrolobatus zone des obersten Karn mit Conodonten-Faunen aus der unteren primitia zone und Halobia septentrionalis ein. Nach einer Lücke im Fossilbefund von 52 m folgt die 200 m mächtige Kerri zone des untersten Nor, welche im unteren Teil Conodonten der oberen primitia zone gemeinsam mit H. cf. beyrichi und H. cf. selwyni erbrachte. Das Ichthyosaurier führende Intervall, von bisherigen Autoren teils ins Karn und/oder ins Nor eingestuft, ist durch eine ergiebige Tropites-Fauna und das Einsetzen von Halobia charakterisiert und kann damit in das obere Karn eingestuft werden. Alle Modelle, die von verschiedenen Autoren für die Erklärung des außergewöhnlichen Ichthyosaurier-Vorkommens vorgeschlagen wurden, werden diskutiert und um ein zusätzliches Genesemodell für die Haupt-Ichthyosaurier-Bank erweitert. Nach der neuen Hypothese führte wohl eine schädliche Algenblüte zu einem Massensterben in diesem Horizont. Obwohl die Karn/Nor Grenze des Brick Pile Aufschlusses im Bereich des fossilleeren 52- m-Intervalls liegt, wird er zu der kleinen Gruppe von Profilen gezählt, welche wesentlich zur Definition des GSSP der Norischen Stufe beitragen können. Die Korrelation von Brick Pile mit den wichtigsten Karn/Nor-Grenzprofilen von British Columbia und Sizilien wird diskutiert. Verglichen mit Juvavites cove und dem GSSP-Kanditaten Black Bear Ridge in B.C., weist Brick Pile bei den Ammonoideen und Halobien einen etwas stärkeren Tethys-Bezug auf. Ein Vergleich von Brick Pile mit dem zweiten GSSP-Kandidaten Pizzo Mondello (Sizilien, Italien) zeigt die signifikanten Probleme, welche bei der Korrelation tethyaler und nordamerikanischer Biochronologien auftreten. Keywords (separated by '-') Upper Triassic - Fossil-Lagerstätte - Nevada - Ammonoids - Halobia - Conodonts - Ichthyosaurs - Mass mortality - Chronostratigraphy Schlüsselwörter (separated Späte Trias - Fossillagerstätte - Nevada - Ammonoideen - Halobia - Conodonten - Ichthyosaurier - by '-') Massensterben - Chronostratigraphie Footnote Information N. J. Silberling: retired. Pala¨ontol Z DOI 10.1007/s12542-014-0244-2

1 RESEARCH PAPER

2 The Carnian/Norian boundary succession at Berlin-Ichthyosaur 3 State Park (Upper Triassic, central NV, USA)

4 Marco Balini • James F. Jenks • Riccardo Martin • 5 Christopher A. McRoberts • Michael J. Orchard • 6 Norman J. Silberling

7 Received: 3 December 2013 / Accepted: 22 August 2014 8 Ó Pala¨ontologische Gesellschaft 2014

Author Proof 9AQ1 Abstract The Upper Carnian-Lower Norian (Upper Tri- Silberling. Halobiids were collected from five levels, and 24 10 assic) Luning Formation at Berlin-Ichthyosaur State Park the first report of conodonts from BISP includes faunas 25 11 (BISP) in central NV (USA) has been sampled using for the from 13 levels. The *340-m thick Brick Pile section, the 26 12 first time the bed-by-bed approach for ammonoids, pelagic most complete in thePROOF study area, includes the uppermost 27 13 bivalves, and conodonts, more than 60 years after its first Carnian Macrolobatus zone, which provides conodont 28 14 description by Silberling (U.S. Geological Survey Profes- faunas of the lower primitia zone and Halobia septentrio- 29 15 sional Paper 322: 1–63, 1959). BISP is historically nalis. The 200-m thick lowermost Norian Kerri zone, 30 16AQ2 important for the definition of the uppermost Carnian of the which begins 52 m above the Macrolobatus zone, yields 31 17 North American Triassic chronostratigraphic scale and is conodonts of the upper primitia zone in its lower part, 32 18 known worldwide as one of the most important ichthyosaur together with H. cf. beyrichi and H. cf. selwyni. The ich- 33 19 fossil-Lagersta¨tte because of its extraordinary record of 37 thyosaur-bearingED interval, whose stratigraphic position has 34 20 articulated, large-sized specimens of Shonisaurus popu- been interpreted quite differently by previous authors, is 35 21 laris. Nearly 190 ammonoids were collected from two documented in the uppermost Carnian Macrolobatus zone 36 22 stratigraphic sections, documenting all the latest Carnian to and is characterized by rich Tropites-dominated ammonoid 37 23 Early Norian ammonoid faunas previously described by faunas and by the onset of Halobia. All models proposed 38 by various workers to explain the unusual ichthyosaur 39 record are discussed and an additional explanation for the 40 main ichthyosaur-bearing bed is proposed. The new 41 hypothesis is that a harmful algal bloom (HAB) may have 42 A1 N. J. Silberling: retired. been the trigger for the mass mortality recorded in this 43 44 & Á level. Although the C/N boundary in the Brick Pile section A2 M. Balini ( ) R. Martin 45 A3 Dipartimento di Scienze della Terra ‘‘Ardito Desio’’, University lies within a 52 m interval that presently lacks paleonto- A4 of Milan, Via Mangiagalli 34, 20133 Milan, Italy logic data, this succession is included in a small group of 46 A5 e-mail: [email protected] sections that are expected to contribute to the definition of 47 the GSSP of the Norian stage. Correlation of the Brick Pile 48 A6 J. F. Jenks 49 A7 1134 Johnson Ridge Lane, West Jordan, UT 84084, USA with the best Carnian/Norian sections in northeastern British Columbia is discussed. Compared to the British 50 A8 C. A. McRoberts Columbia Juvavites cove and the GSSP candidate Black 51 A9 Geology Department, State University of New York at Cortland, Bear Ridge sections, the Brick Pile section exhibits an 52 A10 P.O. Box 2000, Cortland 13045, USA ammonoid and Halobia record that is slightly more similar 53 A11 M. J. Orchard to that of the Tethyan sections. Correlation of the Brick 54 A12 Natural Resources Canada-Geological Survey of Canada, Pile with the second GSSP candidate Pizzo Mondello 55 A13 1500-605 Robson St., Vancouver, BC V6B 5J3, Canada (Sicily, Italy) well demonstrates the significant problems 56 57 A14 N. J. Silberling UNCORRECTencountered in calibration of the Tethyan and North A15 US Geological Survey, Menlo Park, USA American scales. 5859

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60 Keywords Upper Triassic Á Fossil-Lagersta¨tte Á Nevada Á bei der Korrelation tethyaler und nordamerikanischer 111 61 Ammonoids Á Halobia Á Conodonts Á Ichthyosaurs Á Mass Biochronologien auftreten. 112113 62 mortality Á Chronostratigraphy Schlu¨sselwo¨rter Spa¨te Trias Á Fossillagersta¨tte Á 114 63 Kurzfassung Die oberkarnische bis unternorische (Spa¨te Nevada Á Ammonoideen Á Halobia Á Conodonten Á 115 64 Trias) Luning Formation im Berlin-Ichthyosaur State Park Ichthyosaurier Á Massensterben Á Chronostratigraphie 116 65 (BISP) in Zentral-Nevada (USA) wurde mehr als 60 Jahre 66 nach der ersten Beschreibung durch Silberling (U. S. Geo- 67 logical Survey Professional Paper 322: 1–63, 1959) erst- Introduction 117 68 mals bankweise auf Ammoniten, pelagische Bivalven und 69 Conodonten untersucht. Der BISP ist von historischer Be- Berlin-Ichthyosaur State Park (BISP), located in the Sho- AQ3118 70 deutung fu¨r die Definition des obersten Karn in der chro- shone Mountains of central NV (Fig. 1), is a Late Triassic 119 71 nostratigraphischen Tabelle der nordamerikanischen Trias paleontologic locality known worldwide for its unique 120 72 und ist eine der weltweit bekanntesten Ichthyosaurier large-sized ichthyosaur record, consisting of 37 articulated 121 73 Fossillagersta¨tten, ausgezeichnet durch den außergewo¨hn- specimens discovered between the mid-1950s and 1960s 122 74 lichen Fund von 37 artikulierten, großen Exemplaren von within a thin stratigraphic interval of the Luning Formation 123 75 Shonisaurus popularis. Nahezu 190 Ammoniten aus zwei (Camp 1976, 1980). Established in 1957 to protect this 124 76 stratigraphischen Abschnitten belegen oberstes Karn bis exceptional paleontologic locality, BISP’s most impressive 125

Author Proof 77 Unternor, wie schon fru¨her durch Silberling beschrieben. attraction is a sheltered exhibit of several meticulously 126 78 Halobiiden liegen aus fu¨nf Niveaus vor, und der Erst- exposed and well-preserved, *15-m long specimens of the 127 79 nachweis von Conodonten im BISP beinhaltet Faunen aus ichthyosaur Shonisaurus popularis. Indeed, this extraordi- 128 80 13 Niveaus. Der ca. 340 m ma¨chtige Brick Pile Aufschluss nary record of ShonisaurusPROOFled Bottjer et al. (2002) and 129 81 ist der kompletteste im untersuchten Gebiet. Er schließt die Bottjer (2002) to include BISP as one of the most unique 130 82 rund 20 m ma¨chtige Macrolobatus zone des obersten Karn fossil-Lagersta¨tten (Seilacher 1970) sites in the world. 131 83 mit Conodonten-Faunen aus der unteren primitia zone und Notwithstanding the attraction of this Lagersta¨tten to 132 84 Halobia septentrionalis ein. Nach einer Lu¨cke im Fossil- vertebrate paleontologists as well as amateurs, BISP is also 133 85 befund von 52 m folgt die 200 m ma¨chtige Kerri zone des extremely important to invertebrate paleontologists 134 86 untersten Nor, welche im unteren Teil Conodonten der because of the cephalopod, bivalve, and brachiopod faunas 135 87 oberen primitia zone gemeinsam mit H. cf. beyrichi und H. that areED also preserved in the Luning Formation. Its 136 88 cf. selwyni erbrachte. Das Ichthyosaurier fu¨hrende Interv- ammonoid faunas are of outstanding value not only for 137 89 all, von bisherigen Autoren teils ins Karn und/oder ins Nor their abundance and good preservation, but more impor- 138 90 eingestuft, ist durch eine ergiebige Tropites-Fauna und das tantly because they played a crucial role in the definition of 139 91 Einsetzen von Halobia charakterisiert und kann damit in the North American Triassic chronostratigraphic scale 140 92 das obere Karn eingestuft werden. Alle Modelle, die von (Silberling and Tozer 1968; Tozer 1967, 1981b, 1984, 141 93 verschiedenen Autoren fu¨r die Erkla¨rung des au- 1994) that is still regarded as the most complete and finely 142 94 ßergewo¨hnlichen Ichthyosaurier-Vorkommens vorgeschla- subdivided in the world (for historical summary see Tozer AQ4143 95 gen wurden, werden diskutiert und um ein zusa¨tzliches 1984; Balini et al. 2010a, b). 144 96 Genesemodell fu¨r die Haupt-Ichthyosaurier-Bank erweit- Ammonoids were discovered together with the first 145 97 ert. Nach der neuen Hypothese fu¨hrte wohl eine scha¨dliche ichthyosaur remains by Siemon W. Muller of Stanford 146 98 Algenblu¨te zu einem Massensterben in diesem Horizont. University in the late 1920s (Camp 1980: 141–142). 147 99 Obwohl die Karn/Nor Grenze des Brick Pile Aufschlusses Extensive paleontologic investigations were initiated in the 148 100 im Bereich des fossilleeren 52-m-Intervalls liegt, wird er zu early 1950s when Muller’s student, N.J. Silberling, began 149 101 der kleinen Gruppe von Profilen geza¨hlt, welche wesentlich his PhD studies in the Berlin area. In 1953, Muller and 150 102 zur Definition des GSSP der Norischen Stufe beitragen Silberling also guided C.L. Camp, a vertebrate paleontol- 151 103 ko¨nnen. Die Korrelation von Brick Pile mit den wichtigsten ogist from the University of California, Berkeley, in his 152 104 Karn/Nor-Grenzprofilen von British Columbia und Sizilien first survey of the vertebrate localities. Silberling (1959) 153 105 wird diskutiert. Verglichen mit Juvavites cove und dem described the ammonoid faunas in an outstanding mono- 154 106 GSSP-Kanditaten Black Bear Ridge in B.C., weist Brick graph that included not only the first detailed geologic map 155 107 Pile bei den Ammonoideen und Halobien einen etwas of Berlin area, but also the lithostratigraphy of all geologic 156 108 sta¨rkeren Tethys-Bezug auf. Ein Vergleich von Brick Pile formations in the area and the systematic descriptions of 157 109 mit dem zweiten GSSP-Kandidaten Pizzo Mondello ammonoids, nautiloids, and bivalves, as well as the defi- 158 110 (Sizilien, Italien)UNCORRECT zeigt die signifikanten Probleme, welche nition of three ammonoid zones, namely the Klamathies 159

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50 Austin

°30' Wells 9 Winnemucca 121

ange Clan Alpine Mountains R Eastgate Frenchman Bald Mountai n 722

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Toy

Austin '3 Lander Co. Toyabe

39°10 Nye Co. Range 361 Churchill Co. Peak Mineral Co. Tonopah French 376 Peak Sherman Ione Peak G Buffalo Mountai n

abbs Valley ange

R Las Range Gabbs

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N 38°50' Berlin-Ichthyosaur

State Park yabe

20 km To Author Proof 100 Km 118°30' 117°30'

Fig. 1 Location map of Berlin-Ichthyosaur State Park, central NV (USA). A detailed road log with access information for this locality is provided in Lucas et al. (2007) PROOF

Index species Type locality Norian ammonoid-rich successions of British Columbia, 168 169 Juvavites magnus Brown Hill, BC studied by F.H. McLearn between the 1940s and the early 1960s. Thus, correlation of the BISP ammonoid zones with 170 Malayites dawsoni Brown Hill, BC the California successions was proposed by Silberling 171

Lower

NORIAN Stikinoceras kerri Brown Hill, BC (1959), while correlation with those of British Columbia 172 ED 173 Klamathites macrolobatus Shoshone Mts, Nevada was proposed by Silberling and Tozer (1968). The link between the California and British Columbia successions is 174 175 Tropites welleri Shasta County, CA represented by the Macrolobatus zone (type locality BISP,

Upper Shoshone Mountains, NV: Fig. 2), which is bracketed 176 177 Tropites dilleri Shasta County, CA between the underlying Schucherti zone (correlated to the Welleri zone of the Hosselkus Limestone) and the over- 178 CARNIAN Sirenites nanseni Ewe Mountain, BC lying Guembelites zone (correlated to the Lower Norian 179 Austrotrachyceras obesum Ewe Mountain, BC Kerri zone of British Columbia). 180 Lower 181 Trachyceras desatoyense South Canyon, Nevada Despite the world class nature of the fossils preserved in the BISP area, the site apparently has not attracted all that 182 Fig. 2 Lower Carnian-Lower Norian ammonoid zones of the most much attention from paleontologists. Such a conclusion 183 recent version of the North American Triassic chronostratigraphic seems inescapable after an examination of the very scarce 184 scale (Tozer 1994). Zones are represented by index species and type- literature subsequent to Silberling’s work. Since that time, 185 localities (for their definitions, see Tozer 1994). This scale was 186 developed by Tozer and Silberling over a period of about 30 years the locality has been visited by Kristan-Tollmann and (Silberling and Tozer 1968; Tozer 1967, 1981b, 1984, 1994) Tollmann (1983), who illustrated a few Late Carnian and 187 Early Norian ammonoids and halobiids, as well as some 188 160 schucherti, K. macrolobatus, and Guembelites zones in benthic foraminifera. Brachiopods were described by 189 161 ascending chronologic order. Recognition of these zones Sandy and Stanley (1993), and cnidairans, which are rather 190 162 was crucial for the resolution of a bio-chronostratigraphic common in the Luning Formation in the Pilot Mountain 191 163 problem that had hindered the completion of the Upper area (SW of BISP), are known only from a single occur- 192 164 Triassic part of the North American chronostratigraphic rence of a chonrdophorine hydrozoan (Hogler and Hanger 193 165 scale (Fig. 2), i.e., the correlation of the ammonoid faunas 1989). Crustaceans and trace fossils are also known from 194 166 from the Carnian Hosselkus Limestone in northern Cali- single specimens (McMenamin et al. 2013). One would 195 167 fornia, monographedUNCORRECT by Smith (1927), and the Lower expect that such scant invertebrate literature would be 196

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197 greatly overshadowed by literature regarding the area’s Geological and stratigraphic setting 250 198 vertebrate fossils. Surprisingly, this is not the case. There is 199 no record in the literature of any significant new field work Most of BISP is located within West Union Canyon, but a 251 200 in the area since the time of Camp’s extensive excavations. smaller, satellite portion is situated near the mouth of 252 201 Literature published after the 1980s is based either on a re- Berlin Canyon, the next canyon to the north (Fig. 3). These 253 202 examination of Camp’s specimens from a taphonomical two ENE-WSW oriented canyons cut the central western 254 203 point of view (Hogler 1992), or their anatomical restoration slope of the Shoshone Mountains. The ghost-town of 255 204 and taxonomic position (e.g., Kosch 1990; McGowan and Berlin, located in the complex geologic structure of cen- 256 205 Motani 1999). A few authors have attempted to explain the tral-western NV, lends its name to one of the allochthon 257 206 reason for the great number of ichthyosaurs (Massare and units of the Paradise Terrane (sensu Silberling 1991). The 258 207 Callaway 1988; Hogler 1992; McMenamin and Schulte Berlin allochthon (Silberling et al. 1987), together with the 259 208 McMenamin 2011), but no explanation has yet been agreed nearby Lodi allochthon, is characterized as the most 260 209 upon. complete Triassic succession of the Paradise Terrane AQ5261 210 A recent integrated stratigraphic review of continuous (=Paradise subterrane sensu Speed et al. 1989: Fig. 19). 262 211 marine successions pertinent to the definition of the Although the Berlin succession is faulted and thrusted, it is 263 212 GSSP (Global Stratotype Section and Point) of the No- not as deformed and metamorphosed as the Lodi unit. 264 213 rian stage pointed to the potential value of a rigorous The most accurate and complete description of Triassic 265 214 bed-by-bed re-examination of the Carnian-Norian sedi- strata in the BISP area was provided by Silberling, who 266

Author Proof 215 mentary succession in the BISP area. This research, reported a total thickness of about 1.2 km (Silberling 1959: 267 216 carried out within the framework of the activities of the pl. 11; Fig. 4). The Triassic units, lying between the 268 217 Carnian/Norian boundary Working Group of the Sub- Permian Pablo Formation (Ferguson and Cathcart 1954) 269 218 commission on Triassic Stratigraphy, was conceived in and the overlyingPROOF Cenozoic volcanics, consist of the 270 219 order to test the quality of the fossil record in the BISP Middle Triassic Grantsville Formation and the Upper Tri- 271 220 area and to compare it with those of the two best Car- assic Luning Formation. Though highly fractured with 272 221 nian/Norian boundary sections thus far selected as GSSP minor faults, the succession is only cut by two main faults, 273 222 candidates: Black Bear Ridge in British Columbia namely the Richmond Hill Fault (Fig. 3), a west-dipping 274 223 (Canada) and Pizzo Mondello (Sicily, Italy). Compared normal fault, and the West Union Canyon Fault, which 275 224 with these two sections, the ammonoid record of the shows a strike-slip component as demonstrated by the 276 225 BISP succession is known to be richer and more com- positiveED flower structure (Fig. 3) affecting the northern side 277 226 plete, but its halobiid and conodont records are not yet of West Union Canyon. 278 227 that well understood. Conodonts in particular have never Despite the extensive faulting, the stratigraphic rela- 279 228 been studied in the BISP area, and their occurrence tionships of the Permian and Triassic formations are still 280 229 would be important not only to complete the under- well preserved. The Grantsville and Luning formations are 281 230 standing of the BISP fossil record, but also to improve each characterized by a sharp erosional unconformity at 282 231 its correlation with the other two GSSP candidate their base, followed by a transgressive trend that begins 283 232 sections. with coarse-grained conglomerates. The *200-m thick 284 233 In order to test the potential of the BISP succession for Grantsville Formation (Muller and Ferguson 1939) consists 285 234 the definition of the GSSP, a preliminary sampling cam- of a lower clastic member and an upper calcareous mem- 286 235 paign was planned for 2010. Norman J. Silberling was the ber, which yields poorly preserved ammonoids interpreted 287 236 mentor for this project, and he provided invaluable by Silberling (1959) as Ladinian in age. 288 237 assistance during the planning stage of this initial field In the BISP area, the Luning Formation (Muller and 289 238 work. He generously provided us with unpublished data Ferguson 1936, 1939) exhibits a different lithologic suc- 290 239 including his field notes and a stratigraphic log as well as cession with respect to that in its type area (Pilot Moun- 291 240 suggestions for the best sites to sample. Because of his tains, southwestern NV). Silberling (1959) distinguished 292 241 age and deteriorating health, he was unable to participate four informal members in the Shoshone Mountains (Fig. 4) 293 242 in the October 2010 fieldwork, but he maintained contact as follows. The basal clastic member (*200-m thick) is 294 243 with the field team. He also stayed in contact during the overlain by the shaly limestone member (*180 m), which 295 244 initial phase of the paleontologic study of the samples, but includes a 30–40-cm thick alternation of limestones and 296 245 unfortunately he passed away on September 27, 2011. In marls. Comformably overlying the shaly limestone mem- 297 246 honor of his highly significant contribution, we have ber is the *170-m thick calcareous shale member, which 298 247 included Norman in the authorship of this paper and, to is dominated by marls with rare intercalations of thin 299 248 further expressUNCORRECT our gratitude, we dedicate this work to his bedded limestones. The uppermost subdivision is repre- 300 249 memory. sented by the ?450-m thick carbonate member, consisting 301

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2300 Tv 2100 Ppg 2300 Trgc B Tv Gate of Trll the Park Canyon Trgl Berlin Trlc Ppg Trls

Tv Trgc Trgl Trlc BISP boundary Trll Trls Canyon Trlu Trlc Trll A Qw Qw Trll Union 2300 West Union Canyon fault Trlc Author Proof

Ppg West Trlc Trlu PROOF Trls Trll Qw

Trgc 2500

Trlc Qw BISPboundary Trlc Trlc ED Qw Trgl Trgl Tv

Trll

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23 00 Qw Trlc Ppg 1 km 2100 chmond Hill Ri

el Road, gravel road Tv Volcanic rocks livs Trgl Limestone member

t

Contour line (elev. in m) narG

Fm. Trlu Carbonate member Trgc Clastic member Fault, inferred fault noitamroFgni

Stratigraphic section Calcareous shale Greenstone Trls Ppg Ammonoid intervals member . member c: Kerri Zone mFolbaP Shaly limestone b: Macrolobatus Zone Trll Ppl Limestone member a: Schucherti Zone member

n

uL Qw Wash and gravel deposits Trlc Clastic member Ppc Clastic member

Fig. 3 Geological map of West Union Canyon (Shoshone Moun- Silberling (1959). Only two elevation contour lines (2,100 and tains) and Berlin-Ichthyosaur State Park, showing positions of the two 2,300 m) have been reproduced. Star indicates position of the studied stratigraphicUNCORRECT sections (A and B). Map is redrawn from ichthyosaur shelter

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Hogler FormationMember Age (1992) Volcanics

Cenoz. Lithofacies 1200 m Lime mudstones, calcareous shale fossil wacke_ stones with some 1000 carbonate scattered bones Brachiopods stones with secondary dolomite Lime mudstones, bioclastic wacke_ 800 Halobiids Shonisaurus Early Norian Kerri Zone shale

? articulated

calcareous 2 Shonisaurus Limey shales, fossil wackstones laminated mud_ stones and rare

Author Proof 600 Macrolobatus

? sp. Zone Myophoria shoshonensis 1 Studied interval Luning Formation Luning

Schucherti

Septocardia Zone Late Carnian PROOF shaly limestone scattered bones stones and pack_ stones interbedded with clay bioclastic wacke_ rich mudstones 400 A B clastic

erosional unconformity and argillites conglomerate, silty sandstones 200 ED chert pebble lmst. rare ammonoids, bivalves and brachiopods Key for the stratigraphic logs

Ladinian Volcanics Argillite clastic Grantsville Fm. 0 erosional Marl Siltstone unconformity Limestone, Conglomerate dolomite

Fm. Marly Pablo Greenstone Permian limestone

Fig. 4 Triassic lithostratigraphy and fossil record of the Berlin- Shonisaurus remains from nearly the entire Luning Formation, with Ichthyosaur State Park area. Lithostratigraphy and ammonoids from the main concentration in the lowest 50 m of facies equivalent to Silberling (1959), brachiopods from Sandy and Stanley (1993), Silberling’s calcareous shale member (see Hogler 1992: Fig. 3). bivalves from Silberling (1959) and Kristan-Tollmann and Tollmann These two authors also provided different stratigraphic positions for (1983). Two very different interpretations of the stratigraphic the main ichthyosaur bearing level that yielded the specimens distribution of Shonisaurus are presented in the literature. a Camp protected by the on-site shelter (‘‘Fossil House’’ of Hogler 1992): (1980) reported Shonisaurus from Silberling’s shaly limestone (1) position by Camp (1980; see also Fig. 3), (2) position by Hogler member (p. 142–143; see also text); b Hogler (1992, Fig. 3) reported (1992: Fig. 3)

302 of massive limestones with intercalations of thick to very articulated specimens were referred to a 9-m thick interval 310 303 thick beds of crystalline dolomite in its lower part. in the upper part of the member (Camp 1980: 143). The 311 304 The Luning Formation contains an abundant fossil shaly limestone member also provided interesting bivalves 312 305 record, especially in the shaly limestone and calcareous described by Silberling (1959)asSeptocardia? sp. and 313 306 shale members. The shaly limestone member yielded the Myophoria shoshonensis, and more importantly, he repor- 314 307 37 specimens of Shonisaurus collected and described by ted the occurrence of Halobia sp. (op. cit., p. 19) from its 315 308 Camp (1976, 1980). Ichthyousaur remains were reported upper part, although the taxon was not described. The more 316 309 by Camp fromUNCORRECT throughout this member, but many of the common invertebrates of this member, however, are 317

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318 ammonoids and in particular, the Upper Carnian faunas of National Forest (Fig. 3). Areas within BISP are protected, 368 319 the Schucherti and Macrolobatus zones (Fig. 4). Although and it was necessary to secure special permission from the 369 320 Silberling (1959) did not include a range chart for the NV Division of State Parks (NDSP) to conduct paleonto- 370 321 ammonoids found in his bed-by-bed sampled section on the logic sampling during the 2010 field season. The relatively 371 322 southern slope of the West Union Canyon (USGS localities minor portion of field work planned for Toyabe National 372 323 M71 and M72a–c, northwest of the Richmond Mine), he Forest (USFS land) consisted only of surface collecting and 373 324 did report the stratigraphic position of his faunas and in his a special permit was not required for this activity. Actual 374 325 synthetic log (op. cit., p. 37; pl. 11). He also indicated the field work conducted in October 2010 (MB, JJ, and RM) 375 326 position of his ammonoid zones along strike on his included a preliminary survey of both sides of West Union 376 327 1:24.000 scale geological map (Silberling 1959, pl. 10; Canyon and the measurement of two stratigraphic sections. 377 328 Fig. 3). Silberling recognized the Schucherti zone in a These sections were measured with a 20-m long tape and 378 329 38-m thick interval starting about 38 m above the base of compass, and then sampled following a strict bed-by-bed 379 330 the member. He recognized the Macrolobatus zone as approach for ammonoids, bivalves, and conodonts. How- 380 331 extending from about 30 m below the top of the shaly ever, because of permit restrictions, no excavations were 381 332 limestone member (about 76 m above the Schucherti zone) made and samples were taken only from the surface. Very 382 333 through the lowermost 9 m of the overlying calcareous strict procedures were followed regarding sample num- 383 334 shale member. bering and designation. Samples taken from the same level, 384 335 In addition to ammonoids of the uppermost part of the but from different positions along strike were given sepa- 385

Author Proof 336 Macrolobatus zone, the calcareous shale member also rate numbers. The same distinction was applied to float 386 337 includes ammonoid faunas referred to the Kerri zone by pieces of rock moved slightly downslope from their ori- 387 338 Silberling and Tozer (1968). These faunas were reported ginal bed: the eight samples BIS10, BIS10A to 10F, and 388 339 from about 91 to 122 m (USGS Locality M73) above the BIS11 each representPROOF a small, float limestone block col- 389 340 base of the *170-m thick member, i.e., more than 80 m lected no more than 50 cm below limestone bed BIS12. 390 341 above the highest occurring Carnian ammonoid taxon. This Faunal composition, lithology, and thickness of the blocks 391 342 member also yielded halobiids, although very few data are fully support the attribution of these samples to bed BIS12. 392 343 available. Silberling (1959, p. 19) mentioned the presence No ichthyosaur remains were discovered during field 393 344 of Halobia sp., and Kristan-Tollmann and Tollmann (1983) work, but Camp’s Quarry 2 was visited and its location 394 345 figured Perihalobia beyrichi and Halobia hochstetteri from verified on the geological map provided by Silberling 395 346 a locality on the north side of West Union Canyon. (1959ED: Fig. 3). Conodont samples were sent to MJO for 396 347 The carbonate member, the uppermost unit of the Lu- preparation and study, while preparation of macrofossils 397 348 ning Formation in the BISP area, has yielded a few benthic was carried out at the laboratories of the Dipartimento di 398 349 foraminifera (Kristan-Tollmann and Tollmann 1983) and Scienze della Terra (University of Milano). Bivalves were 399 350 brachiopods (Sandy and Stanley 1993), including the spi- sent to CAM for study. All figured fossils are stored at the 400 351 riferid Spondylospira lewesensis (Lees) and the terebratu- New Mexico Museum of Natural History and Science, 401 352 lids Plectoconcha aequiplicata (Gabb), Rhaetina gregaria 1801 Mountain Road NW, Albuquerque, NM 87104-1375. 402 353 Suess, and Zeilleria cf. Z. elliptica. Despite the shallow Inventory numbers are NMMNH P- 67692 to 67737. 403 354 water paleoenvironment, this member has not yet provided 355 cnidarian faunas, even though they are common in the 356 patch reefs of the lower part of the Luning Formation in the The stratigraphic sections 404 357 Pilot Mountain range, close to Mina, NV (Stanley 1977, 358 1979; Kristan-Tollmann and Tollmann 1983; Martindale Exposures of the shaly limestone and calcareous shale 405 359 et al. 2012; Roniewicz and Stanley 2013). members in West Union Canyon are quite poor because of 406 their soft weathering nature and the relatively gentle 407 topography in the outcrop area as well as the heavy veg- 408 360 Materials and methods etation cover (sagebrush, trees, etc.). Nevertheless, Sil- 409 berling’s accurate geological map and the relative 410 361 The stratigraphic interval selected for study includes the abundance of fossil invertebrates made it fairly easy to 411 362 upper part of the shaly limestone member, recording the identify the ammonoid-bearing intervals in the field. Fol- 412 363 uppermost Carnian Macrolobatus zone, and the overlying lowing our survey of both sides of the canyon, the two best 413 364 calcareous shale member, documenting the lowermost exposures were identified; the first (Brick Pile section) is 414 365 Norian Kerri zone. Based on detailed information provided located on the southern side, about 2 km from the mouth of 415 366 by NJS, the areaUNCORRECT chosen for investigation is located in the the canyon, starting from the prominent brick pile in the 416 367 northeastern part of BISP and in the surrounding Toyabe bottom of the canyon (A, Fig. 3), and the second (North 417

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418 section) is located about 2.6 km from the mouth of the Fig. 5 Stratigraphic log of the Luning Formation, Brick Pile section c 419 canyon on its northern side (B, Fig. 3). (West Union Canyon) showing distribution of ammonoids, bivalves 420 (Halobia) and conodonts. Closed circle are highly confident identi- The most complete section (Brick Pile: A, Fig. 3), fications; open circles are poorly preserved specimens 421 measured up-slope to the east from the Brick Pile (WGS84 422 coordinates of the base 38°52041.500N, 117°34058.900W), 423 includes the interval from the upper part of the shaly position within the calcareous shale member. The lower 471 424 limestone member to the base of the carbonate member boundary of the member is covered, and the interval’s 472 425 (Fig. 5), for a total thickness of about 340 m. This partic- position with respect to the lower part of the overlying 473 426 ular section was measured at approximately the same carbonate member cannot be used to estimate its strati- 474 427 location from which Silberling (1959, pl. 11) described the graphic position because of the presence of volcanic rock 475 428 upper portion of his stratigraphic log of the Luning For- outcrops (not shown on the map). The occurrence of three 476 429 mation and subsequently recognized four fossiliferous thick limestone intervals (1.4-, 2-, and 3-m thick: Fig. 6) 477 430 levels (M72a, M72b, M72c, and M73), which correspond suggests an approximate correlation of this section with the 478 431 to the lower, middle, and upper parts of the Macrolobatus upper part of the Brick Pile section, where similar calcar- 479 432 zone and the overlying Guembelites (=Kerri) zone (1959, eous intervals are documented. A more accurate correlation 480 433 p. 37). Site 2 of Kristan-Tollmann and Tollmann (1983: is not possible at this time. 481 434 Fig. 7) also represents the same locality. 435 About 38 m of the Brick Pile section belongs to the

Author Proof 436 shaly limestone member, whereas about 300 m are ascribed Ichthyosaur record 482 437 to the calcareous shale member. This thickness is greater 438 than Silberling’s (1959) original estimate of about 550 ft Although ichthyosaurs cannot serve as a biostratigraphic 483 439 (*167 m) for this member as well as that by Hogler (1992) tool for the definitionPROOF of the Carnian/Norian boundary, it is 484 440 who reported 200 m. Measurements were taken with tape impossible to deal with the BISP fossil record without 485 441 and compass, and bedding measurements were made every referring to this important group of vertebrates. The BISP 486 442 5–10 m. It is highly probable that our measurements are ichthyosaur record, with its numerous articulated and well- 487 443 more accurate than those reported in the literature because preserved individuals, is so remarkable that Bottjer et al. 488 444 the wide variation (21°–52°) in dip angle of the relatively (2002) and Bottjer (2002) were motivated to include the 489 445 few exposed beds along the slope east of the Brick Pile has site in an inventory of the most significant fossil-Lagers- 490 446 a significant influence on the conversion of tape measure- ta¨ttenED in the world. However, with regard to this BISP 491 447 ments into stratigraphic distance. Although the numerous record we must emphasize a significant discrepancy in the 492 448 calcite veins in the lower half of the member point to the literature concerning the stratigraphic position of ichthyo- 493 449 presence of several concealed faults, they most probably do saur remains within the Luning Formation. Figure 4 sum- 494 450 not result in significant repetitions of the succession, at marizes the two distribution models thus far proposed. 495 451 least from the base of the member to level BIS40. In Camp (1980: 142–143), who spent at least seven field 496 452 support of this conclusion, it is noted that on the eastern seasons collecting ichthyosaurs at BISP, described ‘‘all 497 453 side of the West Union Canyon, north of the West Union presently known vertebrates’’ from the shaly limestone 498 454 Canyon fault (Fig. 3), the lower and upper boundaries of member of Silberling (1959), with Quarries X and 4 lying 499 455 the calcareous shale member (easily mappable in agree- within the Schucherti zone, and the other specimens 500 456 ment with Silberling 1959: p. 16) follow a topographic (Quarries 1–3, 5–9) within the Macrolobatus zone. Camp 501 457 course that is consistent with a tabular rock body. Folding (1980: 145–146) also provided descriptions of the type of 502 458 appears to occur only in the uppermost 22 m of the cal- bone preservation, as well as the lithology and stratigraphy 503 459 careous shale member (Fig. 5). Such deformation was of the most important quarries that are all consistent with a 504 460 reported by Silberling (1959: 17) as resulting from ‘‘minor position within the shaly limestone member. 505 461 readjustment’’ between the two contrasting rock types. Hogler (1992) improved the knowledge in many 506 462 The North section (WGS84 coordinates of the base: respects regarding the life and death of the BISP Shoni- 507 463 38°53004.200N, 117°34044.400W) lies entirely within the saurus, by providing a well-presented taphonomical ana- 508 464 calcareous shale member at a site where, according to lysis of bones and articulated specimens (especially those 509 465 Silberling (1959), only the Kerri zone is documented. Kerri preserved under the shelter), as well as a paleobiological 510 466 zone ammonoids and halobiids were also collected by analysis of the ichthyosaur-bearing succession. However, 511 467 Kristan-Tollmann and Tollmann from this site (1983: Hogler’s (1992: Figs. 3, 4) illustrated record of ichthyo- 512 468 locality 6). Unfortunately, the 45-m thick ammonoid- saurs is totally different with respect to that described by 513 469 bearing intervalUNCORRECT is surrounded by a thick debris cover that Camp (1980), which leads to a certain amount of confusion 514 470 makes it nearly impossible to pinpoint its stratigraphic regarding the stratigraphic position and age of Shonisaurus. 515

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West Union Canyon - Brick Pile section

sulfurensis cf. sp. indet.

selwyni beyrichi

C.M. 340 m F cf. cf. 117°34'43.4" W 117°34'43.4" GPS Coordinates: 38°52'43.7" N

ropiceltites

ropites nevadanus

Anatropites silberlingi

Arcestidae

Anatropites

Juvavitinae

Tropithisbites densicostatus Tropithisbites Halobia septentrionalis Guembelites jandianus

Stikinoceras kerri

Halobia Total ammonoids Total

Discophyllites ebneri

Halobia

T Klamathites macrolobatus 320 T

300 BIS42 [4]

BIS41 280

260 communisti . 7 . 8 ex gr. ex gr. Author Proof ? sp. nov. 1 ? sp. nov. 240 sp. nov. 1 sp. nov. sp. nov. 3 sp. nov. sp. nov. 6 sp. nov. sp. nov. 4 sp. nov. 5 sp. nov. sp. nov sp. nov. 1 sp. nov. 2 sp. nov. sp. nov

220 PROOF

Acuminitella acuminata

Acuminitella angusta

Primatella asymmetrica

Primatella mersinensis Primatella Primatella sponge spicules

Primatella

Primatella Primatella Primatella conservativa Primatella Parapetella Metapolygnathus

Kraussodontus microgastropods microbivalves foraminiferids

Primatella Primatella Primatella orchardi Ichthyoliths

200

BIS40 BIS39 BIS20 [11] 180 BIS20b BIS38 [6] BIS21 ED F

160

calcareous shale BIS37 BIS19 BIS36 140 BIS34 [1] F BIS35

120 BIS33 [1] BIS32 LUNING FORMATION LUNING F 100 BIS43 [1]

BIS31 BIS30 ? 80

F 60 F BIS17 [16] BIS16 [4] 40 BIS15 [23] BIS14/15 [1] BIS13,13b [4] M BIS12 [52] BIS11 K BIS10 20 J First level with Tropites bivalves conodonts H (Silberling unpub. data) G

D ammonoids B A GPS Coordinates: 0 shaly limestone UNCORRECT38°52'41.5" N; 117°34'58.9" W

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Fig. 6 Stratigraphic log of the West Union Canyon - North section

Luning Formation, North . 1 section (West Union Canyon), . 1 . 2 . 5

showing the distribution of ex

ammonoids, bivalves (Halobia), ? sp. nov sp. sp. ind. ? sp. ind. tropitiformis orchardi and conodonts asymmetrica sp. nov 3 sp. nov. sp. nov sp. nov 6 sp. nov. 7 sp. nov. 8 sp. nov. cf. 50 communisti otal ammonoids Primatella Primatella gr. gr. Primatella Metapolygnathus Kraussodontus Primatella mersinensis Primatella Primatella Primatella Primatella conservativa Primatella Primatella Ichthyoliths Primatella Acuminitella acuminata Guembelites philostrati Stikinoceras kerri Guembelites Guembelites jandianus Noridiscites Styrites Gonionotites Arcestidae Halobia beyrichi T

40 BIS47 [27] BIS5bis [6] BIS5

30

20

10 BIS4 [20]

BIS2

Author Proof 0 BIS45 [18] BIS46 GPS Coordinates: 38°53'04.2" N; 117°34'44.4" W PROOF 516 Although Hogler did not specifically refer to Silberling’s The confirmation of Camp’s attribution demonstrates a 546 517 lithostratigraphy, her subdivision of the Luning Formation Late Carnian age for the BISP Shonisaurus record. This is 547 518 (Hogler 1992: Fig. 3) can be correlated with that of Sil- in contrast with the Late Carnian to Early Norian age 548 519 berling (Fig. 4). Articulated ichthyosaurs, including those suggested by Hogler’s stratigraphic reconstruction. We 549 520 protected under the shelter (Quarry 2 of Camp 1980, cannot exclude the possibility that some bones/specimens 550 521 equivalent to ‘‘Fossil House’’ of Hogler), are referred to a may occur within the calcareous shale member, as this 551 522 lithofacies equivalent to Silberling’s calcareous shale faciesED is very close to the typical ichthyosaur-bearing 552 523 member, whereas isolated bones were reported as collected facies, but we feel safe in rejecting the carbonate member 553 524 from a lithofacies equivalent to the shaly limestone mem- occurrence. It is difficult to understand why Hogler came 554 525 ber and from two different lithofacies that are equivalent to to this conclusion, but her emphasis was on ichthyosaur 555 526 the carbonate member (see Fig. 4 for summary). This latter study rather than the stratigraphic framework. This 556 527 occurrence is especially surprising because thus far, Tri- approach unfortunately is quite common in marine ver- 557 528 assic ichthyosaurs have never been reported from carbon- tebrate paleontology (see Balini and Renesto 2012), but it 558 529 ate platform facies (e.g., see the general overview by leads to the loss of important information regarding the 559 530 Merriam 1908; Callaway and Massare 1989), and instead understanding of the mode of vertebrate life, as well as 560 531 are typical of the inner- to outer-shelf settings (e.g., Sander their age dating. 561 532 2000). 533 We found no ichthyosaur remains during our 2010 field 534 survey, but we did confirm the lithostratigraphic attribution Ammonoid record 562 535 of Camp’s Quarries 1–3 and 5–9, including Quarry 2 536 (‘‘Fossil House’’), to the shaly limestone member. This As mentioned in the Introduction, literature regarding the 563 537 conclusion is based on (a) the direct experience of NJS; study of ammonoids from BISP is limited to Silberling’s 564 538 (b) Silberling’s highly accurate geological map, which we (1959) monograph and an additional contribution in the 565 539 verified in the field; (c) our verification of the position of early 1980s by Kristan-Tollmann and Tollmann (1983), 566 540 the quarries shown on Silberling’s geological map (Fig. 3); who made a short visit to the site during a tour of Triassic 567 541 and (d) the lithofacies of the sites, especially as regards localities in the American West. Silberling provided a 568 542 Quarry 2. The nine specimens in the sheltered exhibit are complete account of the faunas of the upper part of the 569 543 preserved on top of a *30-cm thick limestone bed, and shaly limestone member and the overlying calcareous shale 570 544 such thick limestone beds are very typical of the shaly member as well as a description of the faunas from a 571 545 limestone member.UNCORRECTtaxonomic point of view. He also included a synthetic 572

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573 stratigraphic chart (pl. 11) and proposed a biostratigraphic Table 1 Cephalopod and bivalve taxa identified in the Upper Car- 574 classification of the succession. nian Schucherti and Macrolobatus zones by Silberling (1959) and 575 The faunas of the Schucherti, Macrolobatus, and Kerri Kristan-Tollmann and Tollmann (1983) 576 zones as documented by Silberling, including a few taxa Macrolobatus zone 577 reported by Kristan-Tollmann and Tollmann (1983), are Silberling (1959) 578 shown with the original taxonomic classification in Klamathites macrolobatus n. sp. (30) 579 Tables 1 and 2. Ammonoid taxonomy was not modified by Tropites latimbilicatus n. sp. (12) 580 Silberling and Tozer (1968) and Tozer (1971), who fol- Tropites subquadratus n. sp. (over 200) 581 lowed Silberling (1959) in considering Stikinoceras Tropites crassicostatus n. sp. (60) 582 McLearn, 1930 as junior synonym of Mojsisovicsites Tropites nodosus n. sp. (24) 583 Gemmellaro, 1904. Afterwards, the separation of the two Tropites nevadanus n. sp. (sev. hundred) 584 genera was suggested by Tozer (1981a) and then confirmed Anatropites sp. (*100) 585 in 1994 in his final review of Canadian Triassic ammonoid Tropiceltites? densicostatus n. sp. (18) 586 faunas. In the same monograph, Tozer established the new Juvavites (Anatomites) cf. inflatus Gemmellaro 587 (monotypic) genus Tropithisbites for Tropiceltites? densi- Griesbachites? cf. cornutus Diener 588 costatus Silberling and described the new species Ana- Arcestes sp. 589 tropites silberlingi for Anatropites sp. of Silberling. Clydonautilus sp. (1) Paranautilus sp.

Author Proof 590 New data Proclydonautilus sp. Aulacoceras sp. 591 The relatively short time period allotted for fossil sampling Kristan-Tollmann and Tollmann (1983) 592 (about 5 days) obviously was not sufficient to allow for the Klamatbites macrolobatusPROOFSilberling 593 collection of a large, representative number of macrofossil Anatropites sp. 594 specimens. However, we did collect (bed-by-bed) about Tropites sp. 595 190 well-preserved ammonoids from the studied interval of Schucherti zone 596 the Luning Formation (Fig. 4). These specimens are sig- Silberling (1959) 597 nificant in that they allow us to outline the ammonoid Klamathites schucherti Smith (50) 598 record, which in turn provides a bio-chronostratigraphic Juvavites (Anatomites) cf. J. (A.) elegans Gemmellaro 599 calibration for Halobia and the collected conodont J. (A.)ED spp. 600 samples. Discophyllites ebneri (Mojsisovics) (6) 601 Ammonoids from the shaly limestone member are nor- Arcestes sp. 602 mally found in limestone beds and are preserved three- Germanonautilus kummeli n. sp. (10) 603 dimensionally. Those from the calcareous shale member Phloioceras mulleri n. sp. (1) 604 are found within marls or in thin marly limestone beds and Proclydonautilus sp. 605 are usually deformed by sediment compaction. Their body ‘‘Orthoceras’’ sp. 606 chambers are nearly always collapsed, with major damage Aulacoceras sp. 607 occurring to the ventral area. Myophoria shoshonensis n. sp. (sev. dozen) 608 The frequency of specimens may change from bed to Septocardia sp. (extr. abundant) 609 bed, but specimens are quite often abundant, especially in Pinna sp. 610 the shaly limestone member (e.g., level BIS12 and blocks Kristan-Tollmann and Tollmann (1983) 611 from this level). Ammonoid bearing levels are rarer in the Klamathites sp. 612 calcareous shale member, but some levels yield a fairly Gonionotites sp. ex gr. italicus 613 high number of specimens (e.g., samples BIS15, BIS17, Arcestes 614 BIS47). All of the more common taxa reported by Silber- Gonionotites sp. 615 ling were found (Figs. 5, 6; Tables 1, 2). These taxa allow Michelinoceras 616 us to identify easily the various ammonoid zones because Nautilus 617 ammonoid faunal composition, even at the generic level, Projuvavites 618 changes quite significantly from one zone to the next Terquemia sp. 619 (Tables 1, 2). See text for explanation of the historical changes in the names of the 620 The Macrolobatus zone is recognized within the Brick zones. Taxa from Silberling (1959) include those described in the 621 Pile section (Fig. 5) from level BIS12, about 5.6-m below Systematic Descriptions and those quoted in the text (p.19). The 622 number of specimens collected is shown in parentheses when available. the top of theUNCORRECT shaly limestone member, to level BIS17, Kristan-Tollmann and Tollmann provided citation of taxa in the text (p. 623 about 12 m above the base of the calcareous shale member. 226 and 228) and some illustrations, but no systematic descriptions

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Table 2 Cephalopod and bivalve taxa identified in the Lower Norian Fig. 7 Ammonoids of the Late Carnian Macrolobatus zone, Luning c Kerri zone by Silberling (1959) and Kristan-Tollmann and Tollmann Formation, Berlin-Ichthyosaur State Park, Brick Pile section. (1983) a–i Shaly limestone member; j–k calcareous shale member. a–c Anatropites silberlingi Tozer, 1994: a specimen NMMNH Kerri zone P-67698 (BIS10b-2), a1 lateral view, a2 apertural view; b specimen Silberling (1959) NMMNH P-67699 (BIS11-13), b1 lateral view, b2 ventral view; Mojsisovicsites robustus (McLearn) (15) c specimen NMMNH P-67700 (BIS6-3), lateral view. d Anatropites sp. ind., NMMNH P-67701 (BIS10f-2), d1 lateral view, d2 ventral Mojsisovicsites kerri (McLearn) (30) view. e–i Tropites nevadanus Silberling, 1959: e specimen NMMNH M. cf. crassecostatus Gemmellaro (40) P-67702 (BIS10-8), depressed morphotype (cf. Silberling 1959: pl. 5, Guembelites clavatus (McLearn) (14) Figs. 3, 4 and 6, 7), e1 lateral view, e2 apertural view; f specimen NMMNH P-67703 (BIS10b-5), f1 lateral view, f2 apertural view; Guembelites jandianus Mojsisovics (*80) g specimen NMMNH P-67704 (BIS10a-2), lateral view; h specimen Guembelites philostrati Diener (10) NMMNH P-67705 (BIS10b-1), lateral view; i specimen NMMNH Styrites cf. subniger Mojsisovics P-67706 (BIS10b-4), i1 lateral view, i2 ventral view. j–k Tropithis- Styrites cf. vermetus (Dittmar) bites densicostatus (Silberling, 1959): j specimen NMMNH P-67707 (BIS16-1), lateral view; k specimen NMMNH P-67708 (BIS26-3), k1 Styrites cf. signatus (Dittmar) lateral view, k2 ventral view. All specimens whitened with ammo- Styrites cf. tropitoides Gemmellaro nium chloride. Bar scale 1 cm for all specimens Dimorphites cf. monti-ignei Dittmar Arcestes sp. Paranautilus sp. The small number of specimens available from BIS13 to 640 Author Proof Cosmonautilus cf. pacificus Smith BIS14/15 for the moment does not allow us to delineate 641 Halobia sp. accurately this boundary, but the faunal change is most 642 643 Kristan-Tollmann and Tollmann (1983) likely related to the facies change. Samples BIS13 and 644 Guembelites clavatus (McLearn) fru¨he Form BIS13b yielded twoPROOF specimens each of Anatropites sil- 645 Guembelites clavatus (McLearn) typische Form berlingi and A. cf. sulfurensis, while BIS14/15 yielded only 646 Guembelites jandianus Mojsisovics one Arcestidae. The upper part of the Macrolobatus zone 647 Guembelites philostrati Diener (levels BIS15-17) is dominated by Tropithisbites densico- 648 Stikinoceras kerri (McLearn) status (Fig. 7j–k) with less abundant Anatropites silberlingi Tozer. 649 Stikinoceras robustus (McLearn) Based on our preliminary sampling, the first occurrence 650 Anatropites sp. of EarlyED Norian Kerri zone ammonoids from the Brick Pile 651 Griesbachites sp. section (Fig. 5) is recorded in level BIS43, about 64 m 652 Thisbites sp. above the base of the calcareous shale member and about 653 Perihalobia beyrichi (Mojsisovics) 52 m above the uppermost ammonoid of the Macrolobatus 654 Halobia hochstetteri (Mojsisovics) zone. In this section the Kerri zone ranges upwards for 655 For explanation of data source see caption for Table 1. The number of about 200 m to level BIS42. The genus Guembelites, 656 specimens collected is shown in brackets when available common from BIS43 to BIS42, is represented by nine 657 specimens from a total of 29. Several partly crushed 658 624 Anatropites silberlingi occurs throughout this interval and specimens attributable to G. cf jandianus were collected 659 625 appears to be the most common taxon of the upper part of from levels BIS43 and BIS35, while well-preserved G. 660 626 the Macrolobatus zone. jandianus Mojsisovics occurs from BIS21 to BIS42 661 627 Level BIS12 is an ammonoid-rich level recognized by (Fig. 8a) together with Stikinoceras kerri McLearn. Such a 662 628 Silberling (1959: 19) as a ‘‘1-ft bed of limestone about high stratigraphic occurrence of G. jandianus and S. kerri 663 629 20 ft below the top of the shaly limestone member’’ is new, because Silberling (1959: pl. 11) reported these two 664 630 characterized by the occurrence of Tropites nevadanus. species only in the lower part of the Kerri zone, while the 665 631 The fauna collected from this level and eight float blocks upper part was characterized by the G. clavatus (McLearn) 666 632 (labels BIS10, BIS10A-10F; BIS11) found a short distance and Mojsisovicsites cf. crassecostatus assemblage. 667 633 down slope from BIS12, consists of Tropites nevadanus This high stratigraphic position of the LO of S. kerri is 668 634 Silberling (Fig. 7e–i) and Anatropites silberlingi Tozer also documented in the North section (Fig. 6), where 669 635 (Fig. 7a–c), with rare Klamathites macrolobatus Silberling, Guembelites and Stikinoceras were collected from the 670 636 Discophyllites ebneri Mojsisovics, and Tropiceltites sp. A lowermost to the uppermost fossil bearing level. In addition 671 637 faunal change occurs around the boundary of the shaly to S. kerri (Fig. 8c, d), level BIS4 is especially rich in 672 638 limestone and calcareous shale members that marks the Guembelites (11 specimens out of a total of 20), with G. 673 639 disappearanceUNCORRECT of Tropites and the onset of Tropithisbites. jandianus the most common species and G. philostrati 674

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UNCORRECT

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Fig. 8 Ammonoids of the Early Norian Kerri zone, Luning Forma- kerri McLearn, 1930: PROOFc specimen NMMNH P-67711 (BIS4-16), tion, calcareous shale member, Berlin-Ichthyosaur State Park. a, North section, lateral view; d specimen NMMNH P-67712 (BIS4-17), Guembelites jandianus Mojsisovics, 1896, NMMNH P-67709 North section, d1 lateral view, d2 ventral view; e Styrites cf. (BIS38-3), Brick Pile section, a1 lateral view, a2 ventral view; tropitiformis Mojsisovics, 1893, NMMNH P-67713 (BIS47-26), b Guembelites philostrati Diener, 1923, NMMNH P-67710 (BIS4- North section, e1 lateral view, e2 apertural view. All specimens 12), North section, b1 lateral view, b2 ventral view. c–d Stikinoceras whitened with ammonium chloride. Bar scale 1 cm for all specimens

675 Diener (Fig. 8b) occurring less frequently. The uppermost TropitesED. This conclusion was based on a population 697 676 fossil bearing levels of the section, BIS5bis and BIS47, analysis of bed-by-bed-collected specimens that led 698 677 yielded the most diverse ammonoid fauna of the two him to differentiate and present a summarized strati- 699 678 stratigraphic sections under study. The largest collection graphic succession (p. 19) of species within the 700 679 was made from level BIS47, whose fauna include Styrites Macrolobatus zone, in ascending order as follows 701 680 cf. tropitiformis (Fig. 8e), Guembelites (G. sp. and G. cf. (measurements represent distance below top of shaly 702 681 clavatus), Stikinoceras kerri McLearn and Gonionotites sp. limestone member): T. latiumbilicatus, 60 ft (18.3 m); 703 682 in decreasing order of frequency. Unfortunately, the T. subquadratus, T. crassicostatus, and T. nodosus, 704 683 lithology of these levels is rather marly and specimens are 50 ft (15.2 m); T. nevadaus, 20 ft (6.1 m). Addition- 705 684 quite often crushed by sediment compaction. It is very ally, more detailed information regarding the range of 706 685 difficult to extract medium to large sized specimens that these individual species within the 18-m thick interval 707 686 are sufficiently complete to permit full classification. is provided in the systematics section of Silberling’s 708 monograph. Within this succession of species, T. 709 subquadratus, T. crassicostatus, and T. nevadanus 710 687 Taxonomic remarks display the widest variability, especially with regard to 711 the height/width (H/W) ratio and the degree of coiling. 712 688 Although the purpose of this paper is not to provide tax- Extreme variants range from morphotypes with a low 713 689 onomic descriptions of the BISP ammonoids, some taxo- degree of coiling and H/W ratios \ or \\ 1 to those 714 690 nomic remarks are necessary in order to explain better the with a medium degree of coiling and H/W *1. Such 715 691 range charts and to introduce the following discussion variability is confirmed by newly collected Tropites 716 692 items. Two points are herein presented: (1) the intraspecific nevadanus specimens from the penultimate bed of the 717 693 variability of Tropites, and (2) a few taxonomic problems shaly limestone member (BIS12 and float samples 718 694 regarding Tropithisbites densicostatus Silberling. BIS10, BIS10A to 10F, and BIS11) (Fig. 7f–i). 719 695 1. Variability of Tropites. Silberling (1959) emphasized 2. Classification of Tropithisbites densicostatus. Speci- 720 696 the wideUNCORRECT intraspecific variability of most species of mens classified as Tropithisbites densicostatus 721

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722 (Fig. 7j–k) fully conform to the original description of independent unit equivalent at least to the upper part of the 772 723 the species given by Silberling (1959), but not to the Welleri zone. Additional elements that must be considered 773 724 interpretation of Tozer (1994). He attributed to T. are the FO of Klamathites macrolobatus, which thus far 774 725 densicostatus Silberling some specimens from the Peril seems to be the marker event of the base of the Macro- 775 726 Formation, Queen Charlotte Island (British Columbia) lobatus zone, and the FO of Anatropites, which will be 776 727 with ribs ‘‘extending to the keel’’ (Tozer 1994: 232). discussed below. 777 728 This particular detail does not agree with the ventral In the upper part of the Macrolobatus zone, Silberling 778 729 side of the BISP specimens described by Silberling (1959: 19, pl. 11) emphasized a key level with Tropites 779 730 (1959: 52) that exhibit furrows bordering the keel. nevadanus, the youngest representative of Tropites in the 780 731 Unfortunately, Tozer included this ‘‘ribs extending to Luning Formation, which is ‘‘confined to a 1-ft bed about 781 732 the keel’’ feature of the Queen Charlotte specimens in 20 ft below the top of the shaly limestone member’’. Our 782 733 the diagnosis of Tropithisbites (p. 231), a new mono- sampling confirms the occurrence of T. nevadanus and K. 783 734 typic genus he erected to accommodate Tropiceltites? macrolobatus in level BIS12 at exactly the same position 784 735 desicostatus Silberling, 1959, which Silberling left in a reported by SIlberling, as well as a lack of Tropites above 785 736 doubtful generic position. According to Tozer, Trop- this level. Although this conclusion is not necessarily 786 737 ithisbites is characterized by a ‘‘ventral keel rounded supported by the very few specimens found in the upper- 787 738 smooth, not bordered by sulci’’, but this diagnosis must most bed of the shaly limestone member (samples BIS13 788 739 be emended because the type species of the genus and BIS13b), it certainly is demonstrated by our more 789

Author Proof 740 exhibits furrows separating the termination of the ribs abundant collections from levels BIS15 to BIS17. 790 741 from the smooth keel. The two most important taxa within the upper part of the 791 Macrolobatus zone are Anatropites silberlingi and Tropi- 792 thisbites densicostatus.PROOF A. silberlingi has been found from 793 742 Biostratigraphic remarks level BIS12 to the top of the zone (BIS17), while T. 794 densicostatus has been collected from levels BIS15 through 795 743 Based on our work thus far, the stratigraphic distribution of BIS17. The bio-chronostratigraphic potential of these two 796 744 ammonoids within the Macrolobatus zone closely agrees taxa is quite different. A facies change most likely influ- 797 745 with that documented by Silberling (1959). This zone ences the FO of T. densicostatus since it first occurs just 798 746 probably can be divided into two different units of possible above the boundary of the shaly limestone-calcareous shale 799 747 zonal rank once three key points are clarified by further members.ED Moreover, given that the paleogeographic dis- 800 748 investigation. These include the stratigraphic position of tribution of this species is unknown (see the taxonomic 801 749 Tropites nodosus Silberling, the FO of the index problems), a separation of the uppermost part of the 802 750 Klamathites macrolobatus, and the FO of Anatropites, all Macrolobatus zone on the basis of this particular taxon 803 751 of which would eventually have to be recorded in the lower would have only local significance. 804 752 part of the Macrolobatus zone, an interval not investigated Conversely, the bio-chronostratigraphic significance of 805 753 in the present work. Anatropites silberlingi is much more important, as dem- 806 754 According to Silberling, the lower and middle parts of onstrated by the following three points: (1) the species is 807 755 the Macrolobatus zone (equivalent to levels J to BIS10: relatively facies-independent, since it occurs both above 808 756 Fig. 5) are characterized by a mass occurrence of Tropites and below the boundary of the shaly limestone and cal- 809 757 as well as by Klamathites macrolobatus. Tropites nodosus careous shale members; (2) the taxon is also known from 810 758 SIlberling is one of five new species of Tropites described northeastern British Columbia, Canada (Tozer 1994), i.e., 811 759 by Silberling from this interval, but it is of special signif- from intermediate paleolatitudes; (3) the genus Anatropites 812 760 icance because it is the only species showing two rows of is very common in Tethyan successions, from western 813 761 distinct and well-developed nodes in both the ventrolateral Tethys to Tethys Himalaya (e.g., Krystyn 1974, 1980, 814 762 and umbilical positions. Such a feature suggests the attri- 1982; Krystyn et al. 2002; Balini et al. 2012), where it is 815 763 bution of T. nodosus Silberling to Margaritropites Diener, the index ammonoid of the uppermost Carnian Spinosus 816 764 1916, as already hypothesized by Krystyn (1982: 32). This zone. Indeed, the main reason we have not yet formally 817 765 genus, however, is thus far known only from the Welleri designated the upper part of the Macrolobatus zone as a 818 766 zone in California and Canada (Tozer 1994) and the new subzone based on this taxon is our uncertainty con- 819 767 equivalent Subbullatus zone in the Tethys realm (Krystyn cerning the stratigraphic position of its FO. We have 820 768 1982). If the occurrence of Margaritropites nodosus is documented A. silberlingi in level BIS12, which is the 821 769 eventually confirmed in the lower part of the Macrolobatus lowermost level thus far sampled, and Silberling did not 822 770 zone as reportedUNCORRECT by Silberling, this datum would support mention Anatropites from levels underlying the bed with 823 771 the separation of this part of the Macrolobatus zone as an Tropites nevadanus. On the other hand, he did not 824

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825 document the non-occurrence of this taxon from the lower (Mojsisovics) and Halobia hochstetteri (Mojsisovics). 875 826 part of the Macrolobatus zone. He only reported (p. 51) Although the specimens illustrated by Gruber (pl. 8, 876 827 Anatropites from the upper part of the shaly limestone Figs. 2, 3 in Kristan-Tollmann and Tollmann 1983) are 877 828 member and the lower part of the calcareous shale member. very poorly preserved, the Perihalobia beyrichi (now 878 829 Though 200-m thick, available data do not justify fur- referred to the genus Halobia) likely corresponds to those 879 830 ther subdivision of the Kerri zone. The FO of Guembelites attributed to this taxon from the Kerri zone (see below). 880 831 (BIS43) is recognized below the FO of Stikiniceras kerri However, the specimens attributed to H. hochstetteri by 881 832 (BIS38), but since we only found three ammonoids below Gruber are somewhat more problematical due to preser- 882 833 level BIS38 (Fig. 5), the discrepancy between the two FOs vation issues and are best considered indeterminate at the 883 834 may not be that significant. With regard to the distribution species level. Regardless, both illustrated specimens are 884 835 of ammonoids within the Kerri zone, a few remarks are reported to come from the calcareous shale member from 885 836 necessary, especially for the occurrence of the three species the north side of West Union Canyon and were reported to 886 837 of Guembelites. be closely associated with several ammonoids including 887 838 Guembelites jandianus is the most common species of Stikinoceras and Guembelites indicative of the Kerri zone 888 839 the Kerri zone in the West Union Canyon sections, but only (see ammonoid discussion above). 889 840 two specimens of G. philostrati and one of G. cf. clavatus The current study resulted in much better preserved 890 841 were collected. This proportion is rather consistent with the halobiids that permit delimitation of at least three distinct 891 842 collections reported by Silberling (1959), who described G. halobiid taxa and also allowed for a better resolution of 892

Author Proof 843 jandianus on the basis of 80 specimens, while G. clavatus latest Carnian and earliest Norian strata (Fig. 9). The 893 844 and G. philostrati were represented in his collection by 14 lowest stratigraphic levels containing halobiids occur on 894 845 and 10 specimens, respectively. The stratigraphic distri- the south side of West Union Canyon at the Brick Pile 895 846 bution of these taxa (Fig. 6) is also consistent with the section. Here, HalobiaPROOF septentrionalis Smith occurs in 896 847 distribution illustrated by Silberling in plate 11, in which level BIS13 (Fig. 9a–b) and is assigned to the Macrolob- 897 848 G. jandianus is shown to occur in the same beds with G. atus zone. These specimens are closely related to Halobia 898 849 philostrati, while G. clavatus occurs in younger levels. radiata Gemmellaro in having densely packed and very 899 850 This record differs significantly from that reported from fine radial ribs, but differ in that the ribs are demarcated by 900 851 other localities in North America and, in part, from the a distinct change in course (the growth stop of Campbell 901 852 Tethys. In British Columbia, Tozer (1994) reported G. 1994; McRoberts 2011). Halobia septentrionalis is closely 902 853 clavatus from subzone 1 of the Kerri zone and G. jandianus relatedED to, if not conspecific with, forms regarded as H. cf. 903 854 from subzone 2. These two species are also known from the rugosa reported from several west-Tethyan localities 904 855 Tethyan Realm, where Krystyn (1982) documented a including the uppermost Carnian at the classic Feuerkogel 905 856 phyletic lineage G. clavatus ? G. jandianus ? G. philo- locality Austria and Pizzo Mondello, Sicily (McRoberts 906 857 strati from the 2nd subzone of the Jandianus zone. How- 2011; Levera 2012). However, Halobia septentrionalis 907 858 ever, the range of G. clavatus In the Tethyan Realm mostly differs from true Halobia rugosa Gu¨mbel, which is rele- 908 859 overlaps the range of G. jandianus, while G. philostrati gated to the Lower Carnian, in its outline and because it has 909 860 occurs in slightly younger beds (Krystyn 1982, Fig. 13). finer radial ornament later in ontogeny. Halobia septen- 910 861 Further investigation would be necessary in the BISP trionalis occurs from several North American localities 911 862 section in order to confirm the occurrence of G. clavatus near the Carnian-Norian boundary and most often in the 912 863 and G. philostrati, as well as their separation from G. uppermost Macrolobatus zone (McRoberts 2011). At the 913 864 jandianus on the basis of a population analysis of their Black Bear Ridge section, H. septentrionalis occurs up to 914 865 intraspecific variability. within 2 cm of the FO of H. austriaca and is relegated to 915 the uppermost Macrolobatus zone. 916 Higher up in the Brick Pile section, specimens referred 917 866 Halobiid record to Halobia cf. selwyini McRoberts and Halobia cf. beyrichi 918 (Mojsisovics) occur about 140 m above the base of the 919 867 Although Silberling (1959) noted the presence of Halobia calcareous shale member. Although determinations of 920 868 from both the shaly limestone and calcareous shale mem- these specimens are not definitive due to small sample sizes 921 869 bers of the Luning Formation in west Union Canyon, they (Fig. 9e, f), they are best compared to Halobia beyrichi 922 870 were either not well enough preserved or too few in (Mojsisovics) and Halobia selwyni McRoberts and are 923 871 number to warrant species-level determination or illustra- likely representative of the Kerri zone. Although H. bey- 924 872 tion. More recently, however, Gruber (in Kristan-Tollmann richi is discussed in more detail below, H. selwyni has 925 873 and TollmannUNCORRECT1983) recognized two halobiid taxa from the previously only been known from its type locality at the 926 874 Luning Formation at Union Canyon: Perihalobia beyrichi Black Bear Ridge section at Williston Lake in northeastern 927

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Fig. 9 Late Carnian and Early Norian Halobia from Luning Formation, Berlin-Ichthyosaur State Park. a–b from the Brick Pile section, shaly limestone member; e–f from the same section, calcareous shale member; c–d from the North section, calcareous shale member. a–b, Halobia septentrionalis Smith 1927, NMMNH P-67692 (BIS13-2), left valve exterior; b Halobia septentrionalis Smith 1927, NMMNH P-67693 (BIS13-2), right valve exterior; c Halobia beyrichi (Mojsisovics 1874), NMMNH P-67694 (BIS45), left valve exterior; d Halobia beyrichi (Mojsisovics 1874), left valve, NMMNH P-67695 (BIS2); e Halobia cf. H. beyrichi (Mojsisovics 1874), Author Proof NMMNH P-67696 (BIS38), right valve exterior; f Halobia cf. H. selwyni McRoberts 2011, slab with numerous individuals, NMMNH P-67697 (BIS21). Bar PROOF scale 1 cm for all specimens

ED

928 British Columbia. At Black Bear Ridge, H. selwyni is Specimens from both horizons (Fig. 9c, d) are relatively 936 929 known from a relatively short interval at the base of the well preserved permitting confident assignment to Halobia 937 930 lower Kerri zone (McRoberts 2011). Additional material beyrichi. This taxon is quite important in being a key zonal 938 931 will be needed to confirm the equivalence of these two index of the Lower Norian in eastern Panthalassa and in the 939 932 taxa. western Tethys (see McRoberts 2010). Unfortunately, a 940 933 From the West Union Canyon North section, Halobia proper evaluation of Mojsisovics’ type specimen of H. 941 934 beyrichi (Mojsisovics) occurs from two stratigraphic levels beyrichi from near Bad Aussee in Austria is not possible (it 942 935 BIS2/45 andUNCORRECT BIS47 in the calcareous shale member. cannot be located in the Geologische Bundesanstalt, 943

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944 Vienna, Austria), its illustration (Mojsisovics 1897, pl. 1, Fig. 10 Conodonts from Luning Formation, Berlin-Ichthyosaur Statec 945 Fig. 7) along with topotype specimens provided by L. Park. a1-3, Primatella conservativa Orchard, NMMNH P-67715 946 (BIS45); b1-3, Primatella conservativa Orchard, NMMNH P-67716 Krystyn and collected by the author confirm species (BIS5); c1-3, Primatella sp. nov. 3, NMMNH P-67717 (BIS45); d1-3, 947 assignment. Similar occurrences elsewhere in western Primatella sp. nov. 5, NMMNH P-67718 (BIS47); e1-3, Primatella 948 North America demonstrably show that it co-occurs with mersinensis (Kozur & Moix), NMMNH P-67719 (BIS47); f1-3, 949 the ammonoid Stikinoceras kerri at several localities Primatella sp. nov. 3, NMMNH P-67720 (BIS5); g1-3, Primatella 950 orchardi (Kozur), NMMNH P-67722 (BIS47); h1-3, Primatella sp. including Vancouver Island and Haida Gwaii (Fredrick nov. 7, NMMNH P-67721 (BIS4); i1-3, Primatella sp. nov. 1, 951 Island), British Columbia, and Kuiu Island, southeast AK NMMNH P-67723 (BIS13b); j1, Primatella sp. nov. 6, NMMNH 952 (McRoberts McRoberts 2010, 2011, and unpbl.) and in the P-67724 (BIS45). Bar scale 200 lm for all specimens 953 western Tethys (e.g., De Capoa Bonardi 1984; Levera 954 2012) where it is closely associated with Halobia styriaca 2013), whereas the other three new species make a later 994 955 (Mojsisovics). At Pizzo Mondello, H. beyrichi first occurs appearance in the upper part of the lower primitia zone. 995 956 approximately 4 m above H. austriaca and in close asso- Primatella sp. nov. 2 ranges no higher than the boundary 996 957 ciation with H. stryiaca higher in the section (Levera fauna characterized as the parvus zone (Orchard 2013). A 997 958 2012). collection from slightly higher in the section (level 998 BIS13b), also yielded Acuminitella angusta Orchard 999 (Fig. 11c) and P. sp. nov. 4, both of which also have a last 1000 959 Conodont record appearance near the top of the parvus zone. Notably absent 1001

Author Proof from each of these collections are representatives of typical 1002 960 Conodonts recovered during this investigation are poorly to Carnian conodonts Kraussodontus and Quadralella, which 1003 961 moderately well preserved, but many are broken and all are are common at Black Bear Ridge, but which largely dis- 1004 962 thermally altered with a color alteration index (CAI) of 5 appear around the Carnian-NorianPROOF boundary after which 1005 963 and occasionally 5.5. Some collections (e.g., BIS30, they become strongly subordinate to both Primatella spe- 1006 964 BIS32) also appear to be more recrystallized and elements cies and diminutive representatives of Metapolygnathus 1007 965 commonly have adhering matrix. Nevertheless, 17 taxa are (including the nominate M. parvus) and Parapetella 1008 966 differentiated and their distribution in both sections of (Orchard 2007, Fig. 5). None of the parvus zone indicators 1009 967 West Union Canyon is shown in Figs. 5 and 6. The ages of occur in the present collections, although they are known to 1010 968 these collections are discussed below with reference to the be widespread (e.g., Carter and Orchard 2013), so the 1011 969 succession established at the prospective GSSP for the lowestED three collections at Brick Pile are judged to lie very 1012 970 Carnian-Norian boundary at Black Bear Ridge, British close to the top of the lower primitia zone in the uppermost 1013 971 Columbia, Canada, where preliminary conodont zones Carnian. 1014 972 have been introduced pending the full description of the Additional conodont collections from the Brick Pile 1015 973 fauna (Orchard 2013). Pending the completion of that work section originate from some 50 m higher and above, 1016 974 and stabilization of the nomenclature, the informal nature spanning some 110 m in total. Level BIS30 marks the 1017 975 of the zones or faunal intervals is emphasized in this appearance of Primatella conservativa Orchard (Fig. 10a, 1018 976 account by denoting them in lower case, i.e. lower, upper, b), which has a long range throughout the primitia zones 1019 977 zones. Of the 17 taxa recorded, all but one are known from at Black Bear Ridge; it is accompanied by most of the 1020 978 Black Bear Ridge. Ten remain undescribed and are kept in species identified in the stratigraphically lower collections. 1021 979 open nomenclature. More significantly, both undisputed Acuminitella acumi- 1022 980 In addition to the conodonts, a variety of other micro- nata Orchard (Fig. 11e–g) and P. asymmetrica Orchard 1023 981 fossils were recovered. Although they do not presently (Fig. 11k, j) appear: these two taxa dominate the fauna 1024 982 provide biochronological constraints, these microfossil and are common through the remainder of the sampled 1025 983 associations may have paleoecological implications. Ich- Brick Pile section. Both species first occur near the top of 1026 984 thyoliths are most common, but phosphatised micromol- the lower primitia zone at Black Bear Ridge, but are more 1027 985 luscan steinkerns are occasionally numerous. Siliceous common in the upper primitia zone. Furthermore, both 1028 986 sponge spicules and foraminiferids occur in one sample species also occur in Haida Gwaii (Carter and Orchard 1029 987 (BIS4 = BIS35: Fig. 5). 2013) and may represent useful Norian indicators. A third 1030 988 Two conodont collections from levels BIS11 and BIS12, species appearing in level BIS30 is Primatella sp. nov. 6 1031 989 from about 30 m in the Brick Pile section, contain long (Fig. 10j), which also appears immediately below the 1032 990 ranging Primatella mersinensis (Kozur and Moix), and a parvus fauna and ranges into the upper primitia zone at 1033 991 further three un-named species of that genus (Figs. 5, 10c, Black Bear Ridge. The same first appearance is recorded 1034 992 e, j). At BlackUNCORRECT Bear Ridge, P. mersinensis ranges for Primatella sp. nov. 7 (Fig. 10g), which occurs in the 1035 993 throughout the lower to upper primitia zones (Orchard next higher sample, level BIS32. In overlying levels 1036

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UNCORRECT

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1037 BIS34 and BIS36, the long ranging Primatella sp. nov. 8 Fig. 11 Conodonts from Luning Formation, Berlin-Ichthyosaur Statec 1038 (Fig. 11d) occurs and in the higher of these two beds it is Park. a1-3, Parapetella sp. nov. 1, NMMNH P-67725 (BIS36); b1-3, 1039 Kraussodontus? sp. nov. 1, NMMNH P-67726 (BIS47); c1-3, accompanied by a single specimen of Parapetella sp. nov. Acuminitella angusta Orchard, NMMNH P-67727 (BIS13b); d1-3, 1040 1 (Fig. 11a), the only representative of this genus so far Primatella sp. nov. 8, NMMNH P-67728 (BIS34); e1, Acuminitella 1041 recovered from the Nevadan section: at Black Bear Ridge, acuminata Orchard, NMMNH P-67729 (BIS30); f2-3, Acuminitella 1042 this species has been found ranging into the parvus zone acuminata Orchard, NMMNH P-67730 (BIS36); g1-3, Acuminitella 1043 acuminata Orchard, NMMNH P-67731 (BIS47); h1-3, Metapolygna- but no higher. The highest collection recovered from the thus ex gr. communisti Hayashi, NMMNH P-67732 (BIS40); i1-3, 1044 east section is from level BIS40 and includes both Primatella sp. nov. 2, NMMNH P-67733 (BIS4); j1-3, Primatella sp. 1045 Primatella orchardi (Kozur) (Fig. 10h) and uncommon nov. 2, NMMNH P-67734 (BIS30); k1-3, Primatella asymmetrica 1046 Metapolygnathus ex gr. communisti Hayashi (Fig. 11h), Orchard, NMMNH P-67735 (BIS4); l1-3, Primatella asymmetrica 1047 Orchard, NMMNH P-67736 (BIS34); m1-3, Primatella sp. nov. 4, both of which have a long range through the boundary NMMNH P-67737 (BIS34). Bar scale 200 lm for all specimens 1048 beds at Black Bear Ridge. In total, the collections from 1049 levels BIS30 through BIS40 are regarded as Norian and 1050 are assigned to the upper primitia zone, which is known to section with regard to the definition of the Carnian/Norian 1087 1051 correspond to the Kerri ammonoid Zone in many Cana- boundary. 1088 1052 dian localities (Orchard and Tozer 1997). Notable also is 1053 the absence of Norigondolella in any of the West Union 1054 Canyon samples: this genus is occasionally abundant in Towards a better understanding of the BISP 1089

Author Proof 1055 Canadian Norian sections, although its sporadic appear- Fossil-Lagersta¨tte 1090 1056 ance there has been related to environmental change. If 1057 present, the position of the distinctive boundary parvus Although the research conducted at BISP in 2010 focused 1091 1058 fauna, is likely to be in the unsampled interval below on the study of the Carnian/NorianPROOF boundary interval with 1092 1059 level BIS30. regard to the definition of the Norian GSSP, the resultant 1093 1060 Four collections of conodonts were recovered from the new data for ammonoids, bivalves, and conodonts improve 1094 1061 relatively short North section (Fig. 6). These contain most the understanding of the unusual BISP Fossil-Lagersta¨tte. 1095 1062 of the species identified in the former section, including, in The most impressive paleontological feature of the Lu- 1096 1063 the lowest sample (BIS45), Acuminitella acuminata, ning Formation within BISP consists of the above-men- 1097 1064 Primatella sp. nov. 7 (Fig. 10g), and P. sp. nov. 8 tioned extraordinary record of 37 articulated specimens of 1098 1065 (Fig. 11d); P. asymmetrica occurs in the next higher ShonisaurusEDfound in the shaly limestone member at 10 1099 1066 sample (BIS4; Fig. 11k). Associated taxa are consistent localities or quarries within the park (Camp 1980). This 1100 1067 with an upper primitia zone age, as implied by co-occur- unusually high number of specimens is especially surpris- 1101 1068 ring Kerri Zone ammonoids. A single specimen from near ing, when one considers that almost all of them were dis- 1102 1069 the top of the section (level BIS47) is referred to Krauss- covered on the surface of natural outcrops. Camp (1980: 1103 1070 odontus? sp. nov. 1 (Fig. 11b), a species that is not known 143) described the ichthyosaur record very meticulously. 1104 1071 from Black Bear Ridge. While most of the specimens were collected from several 1105 beds within a 9-m thick interval in the lower part of the 1106 Macrolobatus Zone, some were discovered on the same 1107 1072 Integrated stratigraphy at BISP and its significance bedding plane. Hogler (1992) very carefully studied the 1108 1073 on a local and global scale specimens from Camp’s Quarry 2, now protected by the 1109 shelter (‘‘Fossil House’’), and recognized nine articulated 1110 1074 Ammonoid data from this study are fully consistent with specimens preserved on a 8 9 20-m bedding plane. 1111 1075 that from the literature (Silberling 1959; Kristan-Tollmann The abundance of ichthyosaur specimens preserved on 1112 1076 and Tollmann 1983), and when examined from a bio- the same bedding plane combined with occurrences in 1113 1077 chronostratigraphic point of view, they can be combined several different levels makes it quite challenging to 1114 1078 with bivalve and conodont data to provide a much better achieve a clear understanding of the BISP Fossil-Lagers- 1115 1079 constrained dating of the Luning Formation. In addition to ta¨tte. Because the BISP Shonisaurus specimens include no 1116 1080 this refined dating, the ensuing integrated stratigraphy trace of soft tissue preservation, the site cannot be con- 1117 1081 based on ammonoids, bivalves and conodonts provides sidered a classic obrutionary Lagersta¨tte (Seilacher 1970; 1118 1082 interesting information useful for the discussion of a set of Seilacher and Westphal 1971; Seilacher et al. 1985). On the 1119 1083 geological problems. On a local scale, the integration of other hand, concentration Lagersta¨tten are usually related 1120 1084 data from the different fossil groups results in an improved to event sedimentation (Seilacher 1970; Seilacher and 1121 1085 understandingUNCORRECT of the BISP Fossil-Lagersta¨tte. On a global Westphal 1971), but the limestone beds of the shaly 1122 1086 scale, the BISP succession is now considered to be a key limestone member of the Luning Formation are described 1123

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ED

UNCORRECT

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1124 as fine grained (Camp 1980; Hogler 1992) and do not Significance of the new data 1177 1125 exhibit features of storm or turbidite deposits. Moreover, 1126 BISP ichthyosaurs are articulated, and such preservation is The most common feature of the four models thus far 1178 1127 not consistent with event sedimentation. proposed to explain the Ichthyosaur record of BISP is the 1179 1128 A number of hypotheses have been proposed to near total absence of significance given to invertebrate 1180 1129 explain the unusual ichthyosaur record of BISP. Camp fossils at this locality. Camp (1980) actually referred to the 1181 1130 (1980: 196–197) suggested a marine sedimentary envi- presence of invertebrates in order to prove a marine envi- 1182 1131 ronment similar to a backwater or embayment with ronment for the Luning Formation, but the weight of this 1183 1132 quiet water for the shaly limestone member of the evidence was not all that important because ichthyosaurs, 1184 1133 Luning Formation. This reconstruction was based on the even in the 1970s, were unknown in fresh waters. More- 1185 1134 combination of fine grained sediments with marine over, his model of intertidal environment for the shaly 1186 1135 invertebrates (brachiopods, clam-like bivalves, cephalo- limestone member is not consistent with the abundance of 1187 1136 pods). Within this environmental framework, he ammonoids and the lithology of limestones that led Sil- 1188 1137 hypothesized that Shonisaurus may have foraged in berling (1959: 16) to reconstruct a normal marine envi- 1189 1138 shallow waters during high tide periods, but then on ronment for this member, documented by echinoid spines 1190 1139 occasion became trapped and was unable to escape identified in thin section, with upward trend of reduction of 1191 1140 during low tides. energy, interpreted as probably due to ‘‘increasing of depth 1192 1141 Another reconstruction was suggested by Massare and of water’’. 1193

Author Proof 1142 Callaway (1988) who emphasized the occurrence of at least Hogler (1992) stated, ‘‘there is no evidence of unusual 1194 1143 one ichthyosaur embryo among the BISP specimens. Camp mortality in the rest of the pelagic fauna’’ (p. 115), but this 1195 1144 (1980: 197) documented the presence of this embryo in the conclusion probably resulted from the erroneous recon- 1196 1145 belly region of specimen A, Quarry 5, but did not attach struction of the stratigraphicPROOF position of the ichthyosaur- 1197 1146 much significance to the occurrence. Massare and Call- rich interval (see Ichthyosaur record chapter). Hogler 1198 1147 away revaluated this specimen and suggested that BISP (1992, Fig. 3) attributed the Shonisaurus-rich interval 1199 1148 may have been part of a breeding or birthing area in order (‘‘Fossil House’’) to the lowest 50 m of the calcareous shale 1200 1149 to explain the rich Shonisaurus record. They also remarked member, which according to Silberling (1959) and our bed- 1201 1150 that the specimens were not deposited simultaneously and by-bed data, is the nearly barren part of the succession. 1202 1151 excluded mass mortality. Conversely, our stratigraphic data fully confirm Camp’s 1203 1152 Hogler (1992) studied in detail the preservation of the attributionED of the 9-m thick ichthyosaur-rich interval 1204 1153 specimens collected by Camp and gave special attention (including those preserved in the ‘‘Fossil House’’) to the 1205 1154 to those protected by the ‘‘Fossil House’’ (Camp’s Macrolobatus Zone (1980:143), and to the portion of the 1206 1155 Quarry 2). She correctly emphasized that the sedimen- succession with the most abundant record of fossil inver- 1207 1156 tological features of the Luning Formation are not con- tebrates. Figure 12, which presents a synthesis of available 1208 1157 sistent with a shallow-water, tide-dominated environment information, includes Silberling’s (1959) data for the 1209 1158 as proposed by Camp (1980), but instead, are indicative Schucherti Zone and lower part of the Macrolobatus Zone 1210 1159 of an off-shore environment. She discussed a combina- as well as our new data from level BIS12 upward. 1211 1160 tion of attritional and catastrophic mortality that could The ichthyosaur-bearing interval not only contains a 1212 1161 account for the scattered isolated specimens and the peculiar ammonoid fauna, but it also records important 1213 1162 closely-spaced large-sized individuals preserved on the faunal changes with regard to bivalves. Septocardia and 1214 1163 same bedding plane (e.g., ‘‘Fossil House’’), respectively. Myophoria, which are abundant in the Schucherti Zone and 1215 1164 However, no particular cause was proffered as most common in the overlying beds without ammonoids, are 1216 1165 likely responsible. replaced by Halobia in the Macrolobatus Zone. The base of 1217 1166 The most recent explanation for the BISP ichthyosaur the Macrolobatus Zone is marked by the onset of Tropites, 1218 1167 record was proposed by McMenamin and Schulte which is extremely abundant from the lower part of this 1219 1168 McMenamin (2011), who bizarrely speculated that ‘‘the zone (see Table 1) through level BIS12. This onset 1220 1169 shonisaurs were killed and carried to the site by an enor- apparently represents a very important environmental 1221 1170 mous Triassic cephalopod, a ‘‘kraken,’’ with an estimated change, since Tropites is a genus with worldwide distri- 1222 1171 length of approximately 30 m, twice that of the modern bution that is often found in high diversity assemblages. 1223 1172 Colossal Squid Mesonychoteuthis’’. However, such a large- Furthermore, Tropites is very common in the Upper Car- 1224 1173 sized cephalopod has never been described, neither for the nian Dilleri and Welleri zones of the North American scale 1225 1174 Triassic, nor for the rest of the evolutionary history of and the Dilleri and Subbullatus zones of the Tethyan scale. 1226 1175 cephalopods,UNCORRECT and there is no science that supports this Indeed, Tropites serves as the index ammonoid for all these 1227 1176 hypothesis. zones, but at BISP this genus does not occur in the 1228

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Berlin-Ichthyosaur State Park Ammonoids

Bivalves Canada Tethys Tozer 1994 Krystyn 1980, 1982 Zones carbonate mb Halobia Guembelites Stikinoceras

2

Kerri Selectus Kerri Lac 1 L. Norian 1 L. Norian Jandianus Noricus Tropites Anatropites Anatropites Tropites Juvavitinae K. macrolobatus Tropites K. schucherti Myophoria Septocardia Margaritropites Anatropites Tropithisbites Shonisaurus calcareous shale mb

BIS12 ? Macrolobatus Macrolobatus Spinosus ? ? 3 Tuv ?

2 Schucherti Welleri Subbullatus shaly limestone mb

1 2 Tuv Upper Carnian Author Proof Upper Carnian (Tuvalian) Rare Ichthyosaur record Common Dilleri Dilleri

poor calibration 1 Tuv Abundant maximum frequency PROOF

Fig. 12 Summary of faunal changes recorded in the shaly limestone correlation with ammonoid successions in British Columbia (Macro- and calcareous shale members of the Luning Formation at BISP. lobatus zone, sensu Tozer 1994) and Tethys. Further sampling is Ammonoid and bivalve data for the Schucherti zone and the lower necessary to confirm the absence of Anatropites in the lower part of part of the Macrolobatus zone are from Silberling (1959). Data from the BISP succession. Thus far, it may be possible that the scope of the level BIS12 and stratigraphically above are fully consistent with that MacrolobatusED zone in British Columbia as recognized by Tozer reported from the same interval by Silberling (1959). Distribution of (1994) overlaps only the upper part of the zone in its type locality Shonisaurus as reported by Camp (1980). Thick dashed line shows (BISP). See text for more detailed discussion the lowermost occurrence of the ammonoid Anatropites and its

1229 Schucherti Zone, which is coeval with the Welleri Zone Acuminatella while less ornate or inornate Kraussodontus, 1249 1230 according to Silberling and Tozer (1968), but instead has Metapolygnathus, and Quadralella are rare. This is also the 1250 1231 its first occurrence in the overlying Macrolobatus zone. case in Kerri zone collections from Black Bear Ridge and 1251 1232 Faunal diversity in the Tropites-rich interval at BISP is, other sections on Williston Lake, but not in the latest 1252 1233 however, quite low. According to Silberling (1959), only Carnian collections. The faunal turnover illustrated by 1253 1234 one or, at the most, two species of Tropites are documented Orchard (2007) in the Williston Lake sections has no 1254 1235 in the same bed, but they always exhibit very wide intra- obvious paleoecological cause within the succession that 1255 1236 specific variability. Such a low diversity faunal composi- Zonneveld et al. (2010) has interpreted as representing 1256 1237 tion clearly reflects a stressed environment that is believed deposition in a deep marine environment. The conodont 1257 1238 to be related to the rich Shonisaurus record. The onset of faunal turnover at Black Bear Ridge, which is regarded as a 1258 1239 Halobia might be related to disaerobic conditions, which significant evolutionary event, corresponds to a small 1259 1240 are quite consistent with the preservation of fully articu- negative shift in organic d13C signaling a reduction in net 1260 1241 lated ichthyosaurs. primary production (Williford et al. 2007). Elsewhere, a 1261 1242 The composition of the conodont faunas is also of positive excursion in carbonate d13C is identified in the 1262 1243 interest in connection with the environmental scenarios CNB interval at Pizzo Mondello (Muttoni et al. 2014), but 1263 1244 discussed above. At the moment, we do not know the this approximates level T2, a deeper stratigraphic level 1264 1245 characteristics of the microfauna below the highest strata compared with the anomaly at Black Bear Ridge. Mazza 1265 1246 assigned to the Macrolobatus zone (BIS12-BIS17). As et al. (2010) has linked the C isotope excursions to changes 1266 1247 noted above, the conodont faunas from West Union Can- in the ratio of conodont genera in Sicily, but this is not 1267 1248 yon are dominatedUNCORRECT by Primatella and similarly ornate evident in British Columbia. 1268

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1269 Although their taxonomic nomenclature differs, Mazza Nautilus. Literature regarding a possible algal blooming 1322 1270 et al. (2010) argued that Epigondolella (in part Primatella influence on fossil cephalopods is also quite meager, con- 1323 1271 of this report) proliferated in seawater with a lower d13C sisting only of a report of mass mortality of Jurassic 1324 1272 relative to Metapolygnathus, Norigondolella, and Para- coleoidea (Wilby et al. 2004). 1325 1273 gondolella (in part Quadralella of this report), which they 1274 regarded as opportunistic genera. Isotope data are not 1275 currently available from West Union Canyon, so we do not Importance of BISP sections for the definition 1326 1276 know whether the rarity of the inornate conodont group in of the GSSP of the Norian stage 1327 1277 the Macrolobatus zone can be explained by generally low 1278 d13C values. At Black Bear Ridge, the inornate group is The importance of the BISP sections and the Brick Pile 1328 1279 still present in the Macrolobatus zone, but is far less section in particular for the definition of the Carnian/No- 1329 1280 common in the Kerri zone, where Norigondolella is also rian boundary has been enhanced significantly with the 1330 1281 occasionally abundant. The virtual exclusion of those discovery of conodonts in the ammonoid and Halobia- 1331 1282 conodonts in the sampled upper Carnian part of the Brick bearing succession. Thus, the Brick Pile section is now 1332 1283 Pile section, and of Norigondolella in the Norian, is included in a very small group of worldwide stratigraphic 1333 1284 anomalous in terms of Canadian data and may be caused by sections that demonstrate a Late Carnian to Early Norian 1334 1285 environmentally restricted conditions, as is proposed to marine fossil record consisting of more than one taxonomic 1335 1286 explain the low diversity of the subjacent Tropites rich group. This small group includes sections at Feuerkogel in 1336

Author Proof 1287 interval. the Northern Alps (Austria; Krystyn 1973, 1980; Krystyn 1337 1288 As a preliminary conclusion, a review of available data and Gallet 2002; Balini et al. 2012), Black Bear Ridge 1338 1289 from BISP suggests a stressed environment during the together with a few nearby sections in northeastern British 1339 1290 deposition of the ichthyosaur-bearing interval. Most Columbia (Canada; TozerPROOF1967, 1994; Orchard et al. 2001; 1340 1291 intriguing is the relationship between Shonisaurus and McRoberts 2007; Orchard 2007; Zonneveld et al. 2010; 1341 1292 Tropites, but this is impossible to resolve with the available McRoberts 2011; Orchard 2013), and Haida Gwaii, B.C. 1342 1293 data. The abundance of articulated, closely spaced Shoni- (Carter and Orchard 2013), a few sections in the Primorye 1343 1294 saurus specimens (‘‘Fossil House’’) is due to a mass mor- region and Yana Okhotskaya River in northeastern Siberia 1344 1295 tality that may have been induced by algal blooming. (Bychkov 1995; Zakharov 1997), Silicka Brezova in Slo- 1345 1296 Although we do not have supporting evidence, this vakia (Krystyn and Gallet 2002), a few sections in Turkey 1346 1297 hypothesis should at least be considered. Harmful algal (Kavaalani,ED Bo¨lu¨cektasi Tepe, and Erenkolu Mezarlik 2: 1347 1298 blooming (HAB) is regarded as one of the most common Krystyn et al. 2002), and Pizzo Mondello in Sicily (Italy; 1348 1299 natural causes of mortality events of marine vertebrates in Muttoni et al. 2001, 2004; Mazza et al. 2010, 2011, 2012; 1349 1300 modern settings, and a suggestion that it may have also Balini 2012; Levera 2012; Muttoni et al. 2014). 1350 1301 occurred in the past was recently advanced by Pyenson Given the condensed nature of the sections from Feu- 1351 1302 et al. (2014), who described an extraordinary accumulation erkogel and Turkey, and the Boreal Realm attribution 1352 1303 of fossil marine vertebrates from a Late Miocene locality in (sensu Tozer 1981b; Dagys 1988) of the northeastern 1353 1304 Atacama, Chile. Further investigation is necessary at BISP Siberian sections, the Brick Pile section is of great signif- 1354 1305 in order to test this hypothesis, whose weak point may be icance as an ideal ‘‘bridge’’ for correlation between the 1355 1306 the relatively deep water deposition of ichthyosaurs in uncondensed mid-paleolatitude sections of British Colum- 1356 1307 contrast with the supratidal stranding reported for ceta- bia and those from the western Tethys because of its rel- 1357 1308 ceans and fishes, and the monospecific composition of the atively low latitude paleogeographic position on the 1358 1309 BISP vertebrate fauna. western margin of North America. At present, however, the 1359 1310 The abundance of Tropites may be due to trophic rela- high resolution correlation of the Brick Pile section with 1360 1311 tionships with Shonisaurus. Either Shonisaurus was feed- the best sections from British Columbia and the western 1361 1312 ing on schools of Tropites,orTropites may have been a Tethys is influenced by the quite different amount of 1362 1313 scavenger, necrophagously feeding on a low oxygenated available data. The two GSSP candidate sections for the 1363 1314 sea bottom rich in organic matter, or even on Shonisaurus definition of the Norian stage, Black Bear Ridge (British 1364 1315 carcasses. The occurrence of Tropites might also have been Columbia) and Pizzo Mondello, have been under study for 1365 1316 influenced by HAB, but again, this is speculation. Many more than 10 years and have been sampled several times 1366 1317 groups of modern cephalopods (e.g., cuttlefishes, squids, with special attention directed at the C/N boundary inter- 1367 1318 and octopods) are known to accumulate HAB toxins and val. In contrast, the Brick Pile section has been sampled 1368 1319 act as vectors in modern food webs (Robertson et al. 2004; only once (2010), but this sampling is considered adequate 1369 1320 Costa et al. 2005UNCORRECT; Bargu et al. 2008; Monteiro and Costa to at least place the position of the C/N boundary within the 1370 1321 2011; Lopes et al. 2013), but no data are available on living 52-m thick still unsampled interval between samples BIS17 1371

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1372 and BIS43 (Fig. 5). Despite the differing amount of Available ammonoid data are shown in Fig. 13, together 1421 1373 available data, we believe that the correlation of the Brick with halobiid data from McRoberts (2011). Since the tax- 1422 1374 Pile section with the most significant Carnian/Norian onomic study of conodonts is still in progress, Fig. 13 does 1423 1375 boundary sections is a worthwhile and necessary interme- not include the range of conodont taxa, but instead presents 1424 1376 diate step in the complex procedure required for selection only the conodont zonation from Orchard (2013). Ammo- 1425 1377 of the Norian GSSP. noid zonation strictly conform to the definition by Tozer 1426 (1994). 1427 1378 Correlation with British Columbia sections Ammonoids are quite rare at BBR and often the samples 1428 consist only of single specimens, many of which are 1429 1379 Several localities in British Columbia have provided Upper sometimes poorly preserved. Gonionotites has been col- 1430 1380 Carnian to Lower Norian fossil records. The most impor- lected from several levels and is the most common 1431 1381 tant, at least with regard to combined macro and micro- ammonoid across the C/N boundary in this section. The 1432 1382 fossil records, is in the Peace River Valley (Eastern most important taxon for the identification of the Macro- 1433 1383 Cordillera), and others are located in Haida Gwaii (for- lobatus zone is Anatropites, often accompanied by Tropi- 1434 1384 merly the Queen Charlotte Islands), such as, for example, celtites. However, Tropiceltites is not a marker of the 1435 1385 at Huxley Island (Orchard 1991). Macrolobatus zone because it can be also found in the 1436 1386 The Peace River area was first surveyed by McLearn in Kerri zone (e.g., Tozer 1994). The same also applies to 1437 1387 the 1930s and 1940s (McLearn 1960 and literature therein), Thisbites, but this genus has never been reported from the 1438

Author Proof 1388 who discovered several localities and provided the first Welleri zone (cf. Tozer 1994); hence, the occurrence of 1439 1389 description of many ammonoids. Many of McLearn’s Thisbites in the lowermost part of the section is used to 1440 1390 localities, however, consisted of small exposures sur- mark the lower boundary of the Macrolobatus zone. 1441 1391 rounded by vegetation, or float blocks from debris. Then, in Discostyrites ireneanusPROOF, index ammonid of Kerri subz- 1442 1392 the 1960s, Tozer discovered many new localities, and with one 1 has not yet been found at BBR, but the lower 1443 1393 the use of improved sampling methods, he reviewed the boundary of this subzone is placed at the FO of Ptero- 1444 1394 Upper Carnian and defined all Lower Norian ammonoid sirenites, even though the chronostratigraphic significance 1445 1395 zones at Brown Hill (Tozer 1965, 1967). The construction of this event is still not yet fully calibrated (see Balini et al. 1446 1396 of the WAC Bennett Dam in the 1960s and the subsequent 2012). This position falls near the base of the parvus zone. 1447 1397 flooding of much of the Peace River Valley in the 1970s The upper boundary of Kerri subzone 1 is marked by the 1448 1398 inundated many of the historical localities, but at the same onlyED occurrence of Guembelites clavatus (McLearn), a 1449 1399 time created new, well exposed, and easily accessible taxon known only from this subzone in Canada (Tozer 1450 1400 outcrops along the shoreline of Williston Lake. These 1994). Kerri subzone 2 is recognized on the occurrence of 1451 1401 outcrops are perfectly suited for bed-by-bed sampling, Stikinoceras kerri McLearn, a rare species at BBR, whose 1452 1402 which was initiated in the early 1980s. A general review of FO is presently placed about 15 m above the occurrence of 1453 1403 the stratigraphic setting of the most important Carnian/ G. clavatus. Conversely the lower part of the Dawsoni zone 1454 1404 Norian boundary sections is provided by Zonneveld et al. is well documented at BBR because Malayites is very 1455 1405 (2010). Black Bear Ridge and Juvavites Cove are among abundant in three levels, starting about 40 m above the 1456 1406 the best sections for comparison with the BISP Brick Pile. section datum. 1457

1407 Black Bear Ridge Juvavites cove 1458

1408 Among the sections exposed along the shoreline of Will- This section is exposed at Pardonet Hill, on the southern 1459 1409 iston Lake, Black Bear Ridge (BBR) is the most interesting shoreline of Williston Lake opposite Black Bear Ridge. 1460 1410 for the definition of the GSSP of the Norian stage. At this Here, the C/N boundary is affected by a change of facies 1461 1411 locality, the C/N boundary interval occurs within the that occurs at the boundary between the Baldonnel and 1462 1412 Pardonet Formation and is not affected by a change of Pardonet formations. However, the section is of great 1463 1413 facies. Consequently, BBR has been proposed as the GSSP interest for its abundant conodont faunas, which are pres- 1464 1414 candidate section (Orchard 2007; Zonneveld et al. 2010; ently under study by MJO, and its well-preserved ammo- 1465 1415 McRoberts and Krystyn 2011). The section is rich in noid faunas of the Kerri and Dawsoni zones. In this respect, 1466 1416 conodonts (Orchard 2007, 2013) and halobiids (McRoberts Juvavites cove provides the best record of the Kerri zone in 1467 1417 2007, 2011), and it has also yielded a relatively poor, but the Williston Lake area, and it is not coincidental that 1468 1418 important ammonoid fauna (Orchard et al. 2001; Krystyn Tozer (1994) defined the lower subdivision of the Kerri 1469 1419 in Balini et al. 2012; Krystyn pers. comm.) as well as a zone (subzone 1, index Discostyrites ireneanus [McLearn]) 1470 UNCORRECT13 1420 record of organic d C variation (Williford et al. 2007). at this section. Bivalve data are not yet available from this 1471

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West Union Canyon - Brick pile Pardonet Hill - Juvavites Cove

55 Black Bear Ridge 340 m Conodont sample GSC loc. # m F

sp. ?

selwyni beyrichi cf. cf.

320 sp. sp. 50 sp.

sp. maclearni Ammonoid Zone sp.

sp.

Anatropites

Klamathites macrolobatus

Tropiceltites

Halobia

Stikinoceras kerri Halobia septentrionalis

Guembelites jandianus

Tropithisbites densicostatus Tropithisbites

Halobia Tropites nevadanus Tropites sp.

300 BIS42 cf. Conodont Zone Ammonoid Zone/Sbz.

sp.

Griesbachites humi

Griesbachites Guembelites clavatus

Thisbites custi Discostyrites ireneanus

Malayites

Gonionotites scapulatus

Malayites bococki

Griesbachites borealis Griesbachites selwyni Pinacoceras

Pterosirenites auritus Stikinoceras kerri Guembelites jandianus

Tropiceltites columbianus Tropiceltites

Thisbites

Gonionotites rarus Dimorphites pardonetiensis 45 BIS41 98505 Ammonoid Zone 280 Conodont Zone

98506 ropiceltites

Griesbachites

Thisbites

Malayites

Anatropites Stikinoceras kerri

Anatropites T Gonionotites Pterosirenites Guembelites clavatus Halobia septentrionalis Halobia ornatissima

Halobia austriaca

Halobia selwyni Halobia radiata 45 m 260

40 Dawsoni 98896 Bed # Sample # 98507 n 240 quadrata 40

o 98508 34

98897 quadrata 35 33 CM99BBR-6 220 98563 98562 35 98899 32 98510 Conodont Zone tion 200 98511 31 10-OF-BB-03 Kerri 30

a 98900 BIS40 30 BIS39 n BIS20 30 180 BIS20b 29 io BIS38 98901 BIS21 25 at

F 28 Kerri Dawsoni primitia m

160 Formati et 25 27 CM99BBR-5 BIS37 Kerri 2 on

BIS19 upper BIS36 CM99BBR-4

d 26 20 C-307864 140 BIS34 g F o r m

r 25

F BIS35 primitia C-307872 24 CM99BBR-3

in 20

Pa 23 C-307829 98561 primitia C-307826

120 BIS33 upper 22 98560 C-307824 un BIS32 21 CM99BBR-4 15 C-307823 L prv upper 20 F For et C-307815 18 C-307812 100 BIS43 n 15 C-307810 17 C-307809 C-307808 C-307807 16 CM99BBR-2 Author Proof BIS31 15 BIS30 10 98516 C-307863 80 ?

ardo 14b ? 14a CM99BBR-1 primitia P 10 14 13c CM01BBR-3 13b F lower 60 ? 13a CM01BBR-2 13 C-307804 F 5 98515 12a C-307803 BIS17 BIS16 5 C-307801 Macrolobatus BIS15 98514 40 BIS14/15 11c 98513 K.1 10b CM01BBR-1 BIS13,13b lwp 10a BIS12 98512 C-307793 9 8 BIS11 0 prv PROOF7 C-307792 n. sp. Q BIS10 bivalves 6 20 J us 5 01-305869 Macrolobatus 0

ammonoids ms

ammonoids L. F. ammonoids bivalves

0 samueli Baldon. Fm.

Ammonoid bio-chronostratigraphy Abbreviations prv: parvus zone Dawsoni highest occurrence of Kerri Zone ammonoids lwp: lower primitia zone Ammonoid indet. Kerri subzone 2 us: upper samueli zone ms: middle samueli zone Kerri subzone 1 ED Macrolobatus

Fig. 13 Correlation of Brick Pile section (BISP) with Juvavites cove Juvavites cove, in order to show the uppermost Carnian to Lower and Black Bear Ridge sections (northeastern British Columbia, Norian zonation of the Pardonet Formation. Consequently, the C/N Canada). The range chart for Brick Pile reports only the most boundary interval, sampled at this locality with centimetric detail (see significant ammonoids and bivalves, see Fig. 5 for the distribution of Orchard 2007; McRoberts 2011), appears very compressed. The all taxa recognized. The range chart for Juvavites cove is new and Norian part of the Brick Pile section is clearly expanded. Ammonoid results from the re-examination (by MB) of all collections from this zones and subzones are recognized following the definition by Tozer locality housed at the Geological Survey of Canada facilities in (1994), while conodont zones follow the classification by Orchard Vancouver. The range chart for Black Bear Ridge includes also data (2013). For details on the lithology of the British Columbia sections, published by McRoberts (2011) and Krystyn in Balini et al. (2012). see Zonneveld and Orchard (2002) and Zonneveld et al. (2010) The scale for the Black Bear Ridge section is optimized with that of

1472 section. The ammonoid faunal composition of some levels consisting of tens of well-preserved specimens are com- 1485 1473 (=GSC localities) was provided by Tozer (1994: appendix), mon, especially in the lower and middle intervals. 1486 1474 but he included only part of the collections from Juvavites Kerri subzone 1 is recognized from the lowermost 1487 1475 cove. All available specimens from this locality, stored at ammonoid level of the section (GSC locality 98512) up to 1488 1476 Geological Survey of Canada in Vancouver, have been level 98514. The parvus conodont fauna is identified a 1489 1477 studied and classified for this work by MB; hence, the short distance below the ammonoid levels, and upper 1490 1478 ammonoid range chart in Fig. 13 provides the most upda- primitia zone faunas occur in association with the Kerri 1491 1479 ted record of ammonoid faunas from this locality. zone macrofauna. Discostyrites ireneanus (McLearn), the 1492 1480 Ammonoids are quite abundant at Juvavites cove and index ammonoid of this subzone, occurs only in level 1493 1481 their preservation is much better than at BBR. They mostly 98514, but the occurrence of Thisbites custi McLearn in 1494 1482 occur in limestone levels in the lowest 5 m of the Pardonet 98512 and 98513, together with Tropiceltites columbianus 1495 1483 Formation as well as in two intervals about 15–20 m and (McLearn), is sufficient evidence to include these levels in 1496 1484 30–45 m aboveUNCORRECT its base (Fig. 13). Ammonoid faunas subzone 1, because these two species have never been 1497

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1498 reported from the Macrolobatus zone (Tozer 1994). The ammonoid level of the section, while conodonts, including 1549 1499 subzonal attribution of level 98515 is unresolved for the those of the parvus zone, have been found in the under- 1550 1500 moment, because its ammonoid assemblage consists of lying beds. Therefore, Kerri subzone 1 at Juvavites cove 1551 1501 Tropiceltites columbianus (McLearn) and Pterosirenites is probably equivalent to only the upper part of this 1552 1502 auritus Tozer, both of which occur in Kerri subzone 1 and subzone at BBR. 1553 1503 2. Correlation of the Williston Lake sections with the Brick 1554 1504 The base of subzone 2 is presently placed at level 98516, Pile section (Fig. 13) is similarly affected by the disho- 1555 1505 based on the occurrence of a poorly preserved specimen mogeneous quality of their records, but despite these lim- 1556 1506 tentatively attributed to Stikinceras kerri McLearn. Most itations, it is possible to make the following observations: 1557 1507 ammonoids were collected from levels 98900 to 98562, 1. The ammonoid, bivalve and conodont record of the 1558 1508 which provided abundant S. kerri together with very well Brick Pile section is fully consistent with that of the 1559 1509 preserved Dimorphites pardonetiansis McLearn, Guemb- two Williston Lake sections, as discussed in the 1560 1510 elites jandianus Mojsisovics, Griesbachites humi preceding Ammonoid, Bivalve, and Conodont record 1561 1511 (McLearn), and the very common but long ranging, Gon- chapters. 1562 1512 ionotites rarus McLearn. Conodonts from these beds 2. For the most part, the BISP sections exhibit only a few 1563 1513 comprise a lower primitia zone fauna. slight differences with respect to the Williston Lake 1564 1514 Level 98897, which yields Griesbachites borealis To- sections in terms of ammonoid faunal composition, 1565 1515 zer, represents the uppermost horizon of subzone 2, and the which suggest a certain amount of Tethyan similarity. 1566 Author Proof 1516 FO of Malayites bococki (McLearn) at GSC 98896 marks These include: (1) the abundance of Tropitidae in the 1567 1517 the base of the Dawsoni zone. The transition from primitia uppermost Carnian; (2) the absence of Pterosirenites in 1568 1518 to quadrata conodont faunas occurs within this uppermost the Norian; (3) the abundance of Guembelites clavatus 1569 1519 part of the Kerri zone. [known only fromPROOF the 2nd subzone of the Jandianus 1570 1520 Correlation discussion zone in the Tethyan Realm (Krystyn 1980, 1982), and 1571 reported only from Kerri subzone 1 in Canada]; (4) the 1572 1521 The Black Bear Ridge and Juvavites cove sections can be occurrence of G. philostrati, thus far not yet found in 1573 1522 easily correlated on the basis of conodont and ammonoid Canada; and (5) the great abundance of Guembelites 1574 1523 biostratigraphy. Correlation of the top of the samueli zone (often representing up to 80 % of some Tethyan 1575 1524 and the lower boundary of the parvus zone document a assemblages:ED Krystyn 1982: 10; see the Ammonoid 1576 1525 hiatus at Juvavites cove at the boundary between the record chapter) combined with the rare occurrence of 1577 1526 Baldonel and the Pardonet formations, where both n. sp. Q Gonionotites. The occurrence of a Halobia taxon 1578 1527 and lower primitia zones are missing. This unconformity is similar to H. beyrichi is also herein considered as 1579 1528 probably diachronous over short distances because possible evidence of Tethyan influence since H. 1580 1529 ammonoid faunas of the Macrolobatus zone (corresponding beyrichi is known to occur in terranes from Nevada 1581 1530 to the missing conodont zones) were first described by to Alaska, but has never been reported from the 1582 1531 Tozer (1965, 1967: Fig. 10) from the Pardonet Formation Williston Lake area (McRoberts 2011). 1583 1532 on the slope of Pardonet Hill, a few hundreds of meters 3. The discovery of conodont faunas at the Brick Pile 1584 1533 upslope from the present day exposure along the lake equivalent to those of British is potentially of great 1585 1534 shoreline. significance for the solution of the correlation of the 1586 1535 The Kerri zone is thicker at Juvavites cove (*38 m) British Columbia conodont scale (Orchard et al. 2001; 1587 1536 than at BBR (*24–25 m), but it is rather difficult to trace Orchard 2007, 2013), and in particular the parvus 1588 1537 laterally specific macrofossil biohorizons within the Kerri zone, with that of the Tethys (Mazza et al. 2010, 2011, 1589 1538 zone because of the large difference in quality and rich- 2012), because Brick Pile shows a certain degree of 1590 1539 ness of the ammonoid record between the two sections. Tethyan affinity. Additional sampling across the C/N 1591 1540 Since conodont-bearing levels are more frequent in both boundary at the Brick Pile section is necessary in order 1592 1541 sections, we consider the conodont zone boundaries closer to determine the presence of this zone, but the 1593 1542 if not equal to the time lines, especially regarding the underlying and overlying conodont faunas of this short 1594 1543 parvus zone (Fig. 13). The parvus zone overlaps the lower bio-chronostratigraphic unit have been already 1595 1544 part of Kerri subzone 1 at BBR, whereas it underlies Kerri identified. 1596 1545 subzone 1 at Juvavites cove. This anomaly is here inter- 4. Available data suggest that the parvus zone and Kerri 1597 1546 preted as due to facies control (and possibly collection subzone 1 correlate with at least part of the *52-m 1598 1547 failure) at the lower boundary of the Kerri subzone 1 at thick unsampled interval in the lower part of the 1599 1548 Juvavites cove.UNCORRECT This boundary coincides with the first calcareous shale member of the Brick Pile section. 1600

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1601 5. The Kerri zone identified in the BISP sections assemblages are dominated by several genera of Sirenitinae 1651 1602 correlates with subzone 2 of the Kerri zone in its type (Bychkov 1995; Zakharov 1997) accompanied by more 1652 1603 area, northeastern British Columbia. This correlation is rare Pinacoceras, Arcestidae, and Phylloceratina. 1653 1604 based on the common occurrence of Guembelites Correlation of the Brick Pile section with Pizzo Mon- 1654 1605 jandianus and Stikinoceras kerri, but Gonionotites dello (Fig. 14), the second GSSP candidate section for the 1655 1606 rarus (McLearn), index ammonoid of this subzone, has base of the Norian, must be examined and it is useful to 1656 1607 not yet been found in the Brick Pile section. emphasize some problems whose solutions are necessary in 1657 1608 6. Integrated chronostratigraphy suggests that the litho- order to define the calibration of the North American and 1658 1609 logic change from the shale-limestone alternation of Tethyan chronostratigraphic scales. 1659 1610 the shaly limestone member to the shale-dominated The Pizzo Mondello succession consists of well-bedded, 1660 1611 calcareous shale member of the Luning Formation light-colored micritic and cherty limestones of the Scillato 1661 1612 (BISP, Brick Pile) resulted from a sudden increase in Formation, which were deposited in a fully pelagic setting. 1662 1613 sedimentation rate. Data in support of this conclusion Although the succession is very rich in conodonts (Mazza 1663 1614 come from the comparison of the thickness of the et al. 2010, 2011, 2012) and Halobia (Levera 2012), as 1664 1615 ammonoid chronozones. The restored record of the well as radiolarian faunas at certain levels (Nicora et al. 1665 1616 Macrolobatus zone at the Brick Pile section is *30 m, 2007; Balini et al. 2010a, b), its ammonoid record is not 1666 1617 the same order of magnitude as the 15 m record at particularly abundant (Balini et al. 2012). Figure 14 pro- 1667 1618 BBR, while the record of the Kerri zone at the Brick vides a summary of ammonoid and Halobia distribution as 1668

Author Proof 1619 Pile section is 200 m, nearly one order of magnitude well as an updated chronostratigraphic subdivision of the 1669 1620 thicker than the *26 m at Juvavites cove. section based on ammonoid occurrences (Balini et al. 1670 1621 7. For further analysis, we suggest that the time duration 2012), which directly tie the succession to Tethyan 1671 1622 of subzone 1 of the Kerri zone, may have been ammonoid chronozonesPROOF and Halobia species (Levera 1672 1623 significantly shorter than that of subzone 2. This 2012), whose ranges are now rather well calibrated with the 1673 1624 hypothesis would require further study, but the docu- ammonoid chronostratigraphy. This correlation required 1674 1625 mented record of subzone 1 at Juvavites cove is about tens of years of research, which was carried out by Krystyn 1675 1626 1 meter, whereas subzone 2 is almost 24 m. This (e.g., 1973, 1974, 1980, 1982; Krystyn et al. 2002; Krystyn 1676 1627 difference is not an artifact due to the hiatus at the in Balini et al. 2012) in the Northern Alps, Turkey, 1677 1628 contact between the Baldonel and Pardonet formations Himalaya, and Timor. 1678 1629 because this gap is documented at the base of the EvenED though ammonoids are quite rare in the succes- 1679 1630 parvus zone. Most probably the lower part of Kerri sion, many taxa such as Discotropites plinii (Mojsisovics), 1680 1631 subzone 1 is not documented by ammonoids at Microtropites, Anatropites, Dimorphites noricus Balini, 1681 1632 Juvavites cove, but the missing part is probably only Krystyn, Levera & Tripodo, and Dimorphites selectus 1682 1633 1–1.5 m, based on the conodont correlation with BBR. Mojsisovics are chronostratigraphically important. These 1683 1634 However, even in this case the corrected Kerri subzone taxa led to the recognition (Balini et al. 2012) of the Dis- 1684 1635 1 would only be about 2.5-m thick, with respect to the cotropites plinii and Gonionotites italicus subzones of the 1685 1636 24 m of Kerri subzone 2. uppermost Carnian Spinosus zone and the Dimorphites 1686 1637 8. The record at the Brick Pile does not detract from the noricus and D. selectus subzones of the lowermost Norian 1687 1638 possibly short duration of Kerri subzone 1. The Jandianus zone. 1688 1639 observed record of the Kerri Zone at the Brick Pile Subbullatus zone 1689 1640 (=to Kerri subzone 2 of British Columbia) is about 1641 200 m, or *four times thicker than the 52-m thick The lowermost part of the Pizzo Mondello section, which is 1690 1642 interval not yet sampled between levels BIS17 and characterized by the occurrence of Halobia carnica Gruber 1691 1643 BIS43, which would be in part the time-equivalent of and Hyattites, is herein tentatively attributed to the Sub- 1692 1644 subzone 1. bullatus zone (=Welleri zone of the North American scale). 1693 Hyattites provides no support to this assignment, however, 1694 1645 Correlation with Pizzo Mondello and other sections because its stratigraphic position cannot be calibrated (see 1695 Balini et al. 2012 for discussion); hence, attribution to the 1696 1646 At present, the Brick Pile section cannot be directly cor- Subbullatus zone relies only on the presence of H. carnica. 1697 1647 related with sections in the Primorye region and the Yana The calibration of this rare species, on the other hand, is not 1698 1648 Okhotskaya River area because of significant paleobioge- considered to be all that precise because H. carnica is 1699 1649 ographic differences that have resulted in vastly dissimilar known only from the Tuvalian 2 of the Raschberg section 1700 1650 ammonoidUNCORRECT faunas. In these regions, ammonoid (northern Alps; Gruber 1976),which Krystyn (1973, 1982) 1701

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West Union Canyon - Brick Pile Pizzo Mondello . Slump

breccia et al level 143 m ammonoids bivalves Conodonts 2001, 2004 Muttoni 140 340 F PM45

m sp. selwyni beyrichi PM44

PM43 affinis paronai cf. cf. f. af cf. 320 130 PM42 sp. sp. ? sp. PM41 sp. ind.

Anatropites Ammonoid Zone

Tropites nevadanus Tropites Tropiceltites Klamathites macrolobatus Tropithisbites densicostatus Tropithisbites Guembelites jandianus Stikinoceras kerri Halobia septentrionalis Halobia

Halobia PM40 300 BIS42 PM39 Anatropites Dimorphites selectus Microtropites Tropiceltites H. austriaca H. styriaca Dimorphites Metathisbites Projuvavites Gonionotites Pregriesbachites bukowskii Dimorphites noricus Placites H. mediterranea Discophyllites insignis 120 H. beyrichi PM38

BIS41 PM37 280 NA49.1 FNP 166.1 PM36 PM36.1 110 PM35 260 PM34 PM33

NA43 PM32 FNP145 240 100 ? FNP144b PM31 FNP144 PM30 NA42.1 PMAM53 FNP135a NA41 220 PM29 PM28d 90 PM28 PM28a

Author Proof NA35 PM27 PMAM35

200 FORMATION

Kerri PMAM43 PM26 BIS40 Conodont Zone PMAM22bis PM25 BIS39 80 PMAM42 FNP112A BIS20 PM24 180 FNP112 BIS20b PMAM7 BIS38 PM23 BIS21 PMAM54 PMAM49 PROOF F PM22 70 PMAM52A 160 PM21 PMAM52 PMAM36 BIS37 PM20 BIS19 FNP88a BIS36 PM19 140 BIS34 60 FNP80A F BIS35 PM18

PM17

120 BIS33 Upper primitia Zone PM16 BIS32 50 PM15 LUNING FORMATION LUNING F ED PM14 BIS43 PM13 100 ? H. lenticularis H. superba BIS31 BIS30 40 PM12 H. simplex 80 PM11 H. radiata

PM10 rugosa cf. O F PM9 60 30 H. F PM8 Discotropites plinii AT

BIS17 Halobia carnica PM7 PMAM27 BIS16 PMAM17 BIS15 PM6 40 BIS14/15 LL BIS13,13b LPZ 20 PM5 BIS12 PMAM15 BIS11 PM4 BIS10 bivalves ? Hyattites 20 FNP331 J ? PM3 FNP333 Macrolobatus SCI ammonoids 10 ammonoids bivalves 0 PM2

PM1 PM0 North american zones and subzones 0 Tethyan zones and subzones

Dawsoni Paulckei

Kerri subzone 2 Selectus sbz. Jandianus Kerri subzone 1 Noricus sbz.

Macrolobatus Spinosus

Fig. 14 Correlation of Brick Pile section (BISP) with Pizzo Mon- Balini et al. (2012) and Levera (2012); the integrated chronostratig- dello section (western Sicily, Italy). The two sections show a raphy is updated from Balini et al. (2012). Thick correlation lines are completely reversed trend in sedimentary evolution: for the Late not time lines s.s. because the fossil-bearing intervals are between Carnian to Early Norian interval the sedimentation rate was notably intervals without fossils. Dashed correlation lines show two alterna- increasing at Brick Pile while at Pizzo Mondello this rate was notably tive options for the correlation of the Welleri/Macrolobatus and the decreasing. TheUNCORRECT range chart of Pizzo Mondello is based on data from Subbullatus/Spinosus boundaries (see text for discussion)

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1702 attributed to the Subbullatus zone on the basis of ammo- Brick Pile section, based on the occurrence of Guembelites. 1751 1703 noid faunal analysis. This taxon has not been found at Pizzo Mondello, but 1752 1704 Correlation of this portion of the Pizzo Mondello section Guembelites is known to occur only in the D. selectus 1753 1705 with the Brick Pile section is thus far uncertain and will subzone in other important localities in the Tethys Realm, 1754 1706 remain so until further fieldwork determines the FO of such as Feuerkogel, type locality of the subzone (section 1755 1707 Anatropites and the position of Margaritroptes at the Brick F1 W: Krystyn 1980; section F5: Krystyn 1980; Balini 1756 1708 Pile. Basically, the problem is not with the Pizzo Mondello et al. 2012), and Jomsom (Krystyn 1982). 1757 1709 section, but instead, it is related to the correlation of this At the present time, the lower subzone of the Jandianus 1758 1710 part of the Tethyan scale with the North American scale. If zone, index taxon Dimorphites noricus, cannot be corre- 1759 1711 the FO of Anatropites is identified at the base of the lated with the Brick Pile section. D. noricus has not yet 1760 1712 Macrolobatus zone, then the Subbullatus zone of the been reported from North America and Halobia austriaca 1761 1713 Tethys (and Pizzo Mondello) can be correlated with the has not yet been found in the Brick Pile section. Thus, 1762 1714 Schucherti zone at BISP (not exposed at Brick Pile sec- correlation of the D. noricus subzone with Kerri subzone 1 1763 1715 tion). If instead the FO of Anatropites is confirmed at level is as problematical as is that of the base of the Macrolob- 1764 1716 BIS10-BIS12 of the Brick Pile section above the atus zone with the base of the Spinosus zones, which well 1765 1717 bed(s) with Margaritropites, it will then be possible to demonstrates the problem of calibration of the Tethyan 1766 1718 correlate the Subbullatus zone of the Tethys and Pizzo scale with the North American scale. 1767 1719 Mondello with the lower part of Macrolobatus zone of the

Author Proof 1720 North American scale. Paulckei zone 1768

1721 Spinosus zone The upper part of the Pizzo Mondello section, ranging from 1769 levels to PM34.1 toPROOF NA51.1 (Levera 2012), is attributed to 1770 1722 The Spinosus zone is about 55 m thick at Pizzo Mondello thePaulckeizonebasedontheoccurrenceofH. mediterra- 1771 1723 and can be directly correlated with at least the upper part of nea, whose chronostratigraphic position is very well con- 1772 1724 the Macrolobatus zone in the Brick Pile section (levels strained (Krystyn et al. 2002). Unfortunately, the few 1773 1725 BIS10,11,12 to BIS17). This correlation is based on the ammonoids from this interval are not age diagnostic: Disco- 1774 1726 occurrence of Anatropites, which has been collected only phyllites and Placites are long-ranging taxa, still consistent 1775 1727 from the upper part of the Spinosus zone at Pizzo Mondello with the Paulckei zone, while the occurrence of Dimorphites 1776 1728 (levels FNP112 and PMAM22bis). However, in the Tethys sp. isED new for this zone. On the whole, the Paulckei zone of 1777 1729 Realm Anatropites ranges from the D. plinii subzone to the Pizzo Mondello is younger than the calcareous shale member 1778 1730 top of the Spinosus zone (cf. Krystyn 1980, 1982). No data of the Luning Formation at the Brick Pile section. 1779 1731 are available on the occurrence of Anatropites within the as 1732 yet unsampled lower part of the Macrolobatus zone at the 1733 Brick Pile (levels J to BIS10, see above), thus correlation Conclusions 1780 1734 of this part with Pizzo Mondello is still unresolved. Integrated stratigraphic research conducted on the Upper 1781 1735 Jandianus zone Carnian/Lower Norian Luning Formation at BISP has 1782 provided a wealth of new data that are of interest for 1783 1736 The Jandianus zone, which is documented at Pizzo Mon- several paleontologic and stratigraphic problems on a local 1784 1737 dello by the occurrence of Dimorphites noricus and D. as well as a global scale. These results are summarized as 1785 1738 selectus from levels NA42.1 to FNP145, accounts for a follows: 1786 1739 thickness of about 3 m. This chronozone’s scope can be 1740 extended both downward and upward based on of the 1. Field work in the Luning Formation of West Union 1787 1741 distribution of Halobia austriaca Mojsisovics, H. styriaca Canyon identified two sections with significant fossil 1788 1742 Mojsisovics, and H. beyrichi Mojsisovics provided by records that were measured and sampled employing a 1789 1743 Levera (2012), because the scope of these species is pre- careful bed-by-bed approach. The Brick Pile section, 1790 1744 cisely calibrated with the ammonoid scale (cf. Krystyn encompassing the Carnian/Norian boundary, is the 1791 1745 et al. 2002; Balini et al. 2012: Fig. 10). Thus, the resulting most complete while the North section is limited to 1792 1746 thickness of the Jandianus zone is about 12.5 m, extending just the Lower Norian. 1793 1747 from the FO of H. austriaca (FNP135a) to the LO of H. 2. Ammonoids, bivalves and, for the first time, con- 1794 1748 beyrichi (FNP154b). The upper part of this interval, from odonts have been found in the studied sections. 1795 1749 the FO of D.UNCORRECT selectus (FNP 144) to the LO of H. beyrichi 3. Ammonoid and bivalve data indicate that the upper- 1796 1750 (FNP154b), can be correlated with the Kerri zone at the most Carnian Macrolobatus zone ranges from the 1797

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1798 shaly limestone member up into the lowermost Guembelites, the occurrence of the Tethyan species 1851 1799 calcareous shale member, while the lowermost Norian G. philostrati, the stratigraphic position of G. clavatus 1852 1800 Kerri zone occurs higher in the calcareous shale and the rare occurrence of Gonionotites. The occur- 1853 1801 member. Conodont data are fairly consistent with the rence of an undescribed species of Halobia similar to 1854 1802 ammonoid and bivalve record, but they do not yet H. beyrichi may also provide additional evidence of 1855 1803 permit high-resolution dating. Conodont faunas found Tethyan influence. 1856 1804 in the Macrolobatus zone are referred to the lower 9. Correlation of the Brick Pile section with Juvavites 1857 1805 primitia zone, while those from the Kerri zone cove and Black Bear Ridge shows that the thickness 1858 1806 document the upper primitia zone sensu Orchard of the Macrolobatus zone is of the same order of 1859 1807 (2013). magnitude as the British Columbia sections, while the 1860 1808 4. The stratigraphic position of the ichthyosaur-bearing scope of the Kerri zone is at least one order of 1861 1809 interval at BISP is revised. The ichthyosaur interval magnitude thicker. This suggests that the lithologic 1862 1810 occurs within the shaly limestone member of the change from the shaly limestone member to the shale- 1863 1811 Luning Formation in the Upper Carnian Macrolobatus dominated calcareous shale member of the Luning 1864 1812 zone. This interval is characterized by rich Tropites- Formation at the Brick Pile (BISP) resulted from a 1865 1813 dominated ammonoid assemblages and a bivalve huge increase in sedimentation rates. Comparison of 1866 1814 faunal turnover from Septocardia-dominated to Halo- the records of subzone 1 and 2 of the Kerri zone 1867 1815 bia-dominated assemblages. suggest a significantly shorter time-duration for 1868

Author Proof 1816 5. Models suggested by various workers to explain the subzone 1 with respect to subzone 2. 1869 1817 accumulation of large-sized, closely spaced ichthyo- 10. Correlation of the BISP Brick Pile section with Pizzo 1870 1818 saurs such as those preserved in the ‘‘Fossil House’’ Mondello demonstrates the difficulties encountered 1871 1819 are discussed. Most of these have weak points or are when attemptingPROOF to calibrate the Tethyan and North 1872 1820 not consistent with available geologic and paleonto- American chronostratigraphic scales. Significant 1873 1821 logic data, which document a relatively deep, problems include correlation of the boundary between 1874 1822 stressed, and low oxygenated/dysoxic environment the Welleri/Macrolobatus zones with that of the 1875 1823 during the deposition of the ichthyosaur-bearing Subbullatus/Spinosus zones and correlation of the 1876 1824 interval. As pure speculation, we suggest that the boundary between the Macrolobatus/Kerri zones with 1877 1825 accumulation of ichthyosaurs may have been influ- the Spinosus/Jandianus zones. The solution to the first 1878 1826 enced by harmful algal blooming (HAB). problemED requires the location of the FO of Anatrop- 1879 1827 6. Based on our new data, the stratigraphic position and ites by bed-by-bed sampling of the North American 1880 1828 range of the Macrolobatus zone is confirmed as section(s), a bioevent that has already been deter- 1881 1829 reported by Silberling (1959), while the range of the mined for several Tethyan sections. In this respect, 1882 1830 Kerri zone is extended downwards significantly. Its the Brick Pile section appears to be the only key 1883 1831 range is now about 200 m, which is more than twice section in North America. Resolution of the second 1884 1832 that reported by Silberling (1959). significant problem is more difficult because the exact 1885 1833 7. The Upper Carnian-Lower Norian record of the BISP location of the Macrolobatus/Kerri boundary has yet 1886 1834 Brick Pile section is compared with that of the most to be documented within the unsampled 52-m thick 1887 1835 important sections in the world for the C/N boundary interval between the last sample providing Carnian 1888 1836 definition: Black Bear Ridge (GSSP candidate) and fossils and the first level yielding Norian fossils. 1889 1837 Juvavites cove in northeastern British Columbia Most of the unresolved issues resulting from the 1890 1838 (Canada), and Pizzo Mondello (GSSP candidate) in investigation initiated in 2010, are specifically addressed 1891 1839 southern Italy. All available ammonoid collections in a new field plan scheduled for summer 2015, when 1892 1840 from Juvavites cove have been reviewed to ensure the the Brick Pile section will be trenched from the base of 1893 1841 most up-to-date and accurate correlation. Moreover, the Macrolobatus zone to the middle part of the Kerri 1894 1842 an updated ammonoid-bivalve integrated chronostra- zone. 1895 1843 tigraphy of Pizzo Mondello section is provided. 1844 8. The BISP Brick Pile section is easily correlated with Acknowledgments We kindly thank the Nevada Division of State 1896 1845 Black Bear Ridge and Juvavites cove on the basis of Parks and Nevada State Museum personnel who were responsible for 1897 1846 the three taxonomic groups discussed in this work. expediting approval of our special permit application for paleonto- 1898 1847 logic work within BISP. Their efforts are greatly appreciated. Like- 1899 Ammonoid faunal composition suggests a Tethyan 1900 1848 wise, we are appreciative of the US Forest Service for allowing us to influence on the BISP faunas because of the abun- conduct reconnaissance work on Toyable National Forest land. The 1901 1849 dance of Tropitidae in the Carnian, the lack of on-site assistance of BISP supervisor Jeff Morris and his staff during 1902 UNCORRECT 1903 1850 Pterosirenites in the Norian, the abundance of our very rainy October 2010 field trip was also greatly appreciated.

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1904 Ammonoid taxonomic problems were discussed with Leo Krystyn Magadan: Severo-Vostocnyj Nauchnyj Center Dalnevostoch- 1966 1905 (University of Vienna, Austria), who is the acknowledged leading nogo Otdeleniya RAN. (in Russian). 1967 1906 authority on Carnian-Norian ammonoid systematics and biostratig- Callaway, J.M., and J.A. Massare. 1989. Geographic and Strati- 1968 1907 raphy. Many thanks to Hans Hagdorn (Ingelfingen) for the stimulating graphic Distribution of the Triassic Ichthyosauria (Reptilia; 1969 1908 discussions on concept and definition of mass mortality and Fossil- Diapsida). Neues Jahrbuch fu¨r Geologie und Pala¨ontologie 178: 1970 1909 Lagestatten. Spencer Lucas and Justin Spielmann (New Mexico 37–58. 1971 1910 Museum of Natural History and Science, Albuquerque, NM) kindly Camp, C.L. 1976. Vorla¨ufige Mitteilungen u¨ber große Ichthyosaurier 1972 1911 helped with registration of the published specimens in the catalogue aus der oberen Trias von Nevada. O¨ sterreichischen Akademie 1973 1912 of the NMMNHS. Warm thanks also to Alexander Lukeneder (Na- der Wissenschaften, Sitzungsberichte der mathematisch-natur- 1974 1913 turhistorisches Museum, Wien) and Leo Krystyn who kindly trans- wissenschaftlichen Klasse Abt. I 185: 125–134. 1975 1914 lated the Abstract into German. Camp, C.L. 1980. Large Ichthyosaurs from the Upper Triassic of 1976 1915 The manuscript has been notably improved by valuable and stim- Nevada. Palaeontograhica Abt. A 170: 139–200. 1977 1916 ulating suggestions by Leo Krystyn and an anonymous reviewer. Campbell, H.J. 1994. The Triassic bivalves Halobia and Daonella in 1978 1917 Technical support for the ammonoid part was provided by G. New Zealand, New Caledonia, and Svalbard. Institute of 1979 1918 Chiodi (University of Milano). This work is a contribution to PRIN Geological & Nuclear Sciences Monograph 4: 1–166. 1980 1919 2008 project ‘‘Stratigrafia integrata del Triassico Superiore: GSSP e Carter, E.S., and M.J. Orchard. 2013. Intercalibration of conodont 1981 1920 sezioni ausiliarie in Italia’’ (P.I. M. Balini). Financial support for field and radiolarian faunas from the Carnian-Norian Boundary 1982 1921 works has been provided by PRIN 2008 grants to M. Balini (Milano Interval in Haida Gwaii, British Columbia, Canada. In The AQ71983 1922 Research Unit). The conodont analyses were undertaken in Vancou- Triassic system, 61st ed, ed. L.H. Tanner, J.A. Spielman, and 1984 1923 ver as a contribution to the Geological Survey of Canada’s ‘‘Geo- S.G. Lucas, 67–92. Albuquerque: New Mexico Museum of 1985 1924 science for Energy and Minerals’’ (GEM) program, with support from Natural History and Science, Bulletin. 1986 1925 the Yukon Basins Project (M.J. Orchard). Costa, P.R., R. Rosa, J. Pereira, and M.A.M. Sampayo. 2005. 1987 Detection of domoic acid, the amnesic shellfish toxin, in the 1988 Author Proof digestive gland of Eledone cirrhosa and E. moschata (Cepha- 1989 lopoda, Octopoda) from the Portuguese coast. Aquatic Living 1990 1926 References Resources 18: 395–400. 1991 Dagys, A.S. 1988. Major features of the geographic differentiation of 1992 1927AQ6 Arkell, W.J., B. Kummel, and C.W. Wright. 1957. Mesozoic triassic ammonoids.PROOF In Cephalopods present and past, ed. 1993 1928 ammonoidea. In Treatise on invertebrate paleontology, part L, J. Wiedmann, and J. Kullmann, 341–349. Stuttgart: Schweizer- 1994 1929 mollusca 4—cephalopoda, ammonoidea, ed. W.J. Arkell, W.M. bart’sche Verlagsbuchhandlung. 1995 1930 Furnisch, B. Kummel, A.K. Miller, R.C. Moore, O.H. Schinde- De Capoa Bonardi, P. 1984. Halobia zones in the pelagic Late 1996 1931 wolf, P.C. Sylvester Bradley, and C.W. Wright, L80–L490. Triassic sequences of the central Mediterranean area. 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2164 Nichols, K.M., and N.J. Silberling. 1977. Stratigraphy and Deposi- 5–23 Philosophical Transactions of the Royal Society of 2227 2165 tional History of the Star Peak Group (Triassic), Northwestern London B. 2228 2166 Nevada. Geological Society of America Special Paper 178: Silberling, N.J. 1959. Pre-Tertiary Stratigraphy and Upper Triassic 2229 2167 1–73. Paleontology of the Union District Shoshone Mounatins, 2230 2168 Nicora, A., M. Balini, A. Bellanca, A. Bertinelli, S.A. Bowring, P. Di Nevada. U. S. Geological Survey Professional Paper 322: 1–67. 2231 2169 Stefano, P. Dumitrica, C. Guaiumi, M. Gullo, A. Hungerbuehler, Silberling, N.J., and R.J. Roberts. 1962. Pre-Tertiary stratigraphy and 2232 2170 M. Levera, M. Mazza, C.A. McRoberts, G. Muttoni, N. Preto, structure of northwestern Nevada. Geological Society of Amer- 2233 2171 and M. Rigo. 2007. 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Canadian Journal of Earth Sciences 8: 989–1031. 2258 2196 and sequence stratigraphy of the Upper Triassic of the Black Tozer, E.T. 1979. Latest Triassic ammonoid faunas and biochronol- 2259 2197 Bear Ridge, northeast British Columbia, a GSSP prospect for the ogy, Western Canada. Current Research, Part B, Geol. Surv. 2260 2198 Carnian-Norian boundary. Albertiana 25: 10–22. Canada Paper 79-1B: 127–135. 2261 2199 Pyenson, N.D., C.S. Gutstein, J.F. Parham, J.P. Le Roux, C.C. Tozer,ED E.T. 1981a. Triassic Ammonoidea: Classification, Evolution 2262 2200 Chavarrı´a, H. Little, A. Metallo, V. Rossi, A.M. Valenzuela- and Relationship, with Permian and Jurassic Forms. In The 2263 2201 Toro, J. Velez-Juarbe, C.M. Santelli, D. Rubilar Rogers, M.A. Ammonoidea, ed. M.R. House and J.R. Senior, The Systematic 2264 2202 Cozzuol, and M.E. Sua´rez. 2014. Repeated mass strandings of Association special volume 18: 65–100. London: Acadamic 2265 2203 Miocene marine mammals from Atacama Region of Chile point Press. 2266 2204 to sudden death at sea. Proceedings of the Royal Society B: Tozer, E.T. 1981b. Triassic Ammonoidea: Geographic and Strati- 2267 2205 Biological Sciences 281(1781): 20133316. graphic Distribution. In The Ammonoidea, ed. M.R. House and 2268 2206 Robertson, A., D. Stirling, C. Robillot, L. Llewellyn, and A. Negri. J.R. Senior, The Systematic Association special volume 18: 2269 2207 2004. First report of saxitoxin in octopi. Toxicon 44: 765–771. 397–432. London, New York: Acadamic Press. 2270 2208 Roniewicz, E., and G.D. Stanley. 2013. Upper Triassic corals from Tozer, E.T. 1984. The Trias and its ammonoids: the evolution of a 2271 2209 Nevada, western North America, and the implications for time scale. Geological Survey of Canada, Miscellaneous Report 2272 2210 paleoecology and paleogeography. Journal of Paleontology 87: 35: 1–171. 2273 2211 934–964. Tozer, E.T. 1994. 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A record of stable organic carbon isotopes 2281 2219 1970: 34–39. from the Carnian-Norian boundary section at Black Bear Ridge, 2282 2220 Seilacher, A. and F. Westphal. 1971. Fossil-Lagersta¨tten. In Sedi- Williston Lake, British Columbia, Canada. Albertiana 36: 2283 2221 mentology of parts of Central Europe, Guidebook 8. Int. 146–148. 2284 2222 Sediment. Congr. 327–335, Heidelberg. Zakharov, Yu.D. 1997. Carnian and Norian Sirenitid ammonoids of 2285 2223 Seilacher, A., W.E. Reif, and F. Westphal, 1985. Sedimentological, the north-western circum-pacific and their role in the Late 2286 2224 ecological and temporal patterns of fossil Lagersta¨tten. In Triassic faunal succession. In Late Paleozoic and Early Meso- 2287 2225 Extraordinary fossil biotas: their ecological and evolutionary zoic Circum-Pacific Events: Biostratigraphy, Tectonic and Ore 2288 2226 2289 significanceUNCORRECT, ed. H.B. Whittington and S. Conway Morris, 311, Deposits of Primoryie (Far East Russia), ed. A. Baud, I. Popova,

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2290 J.M. Dickins, S. Lucas and Yu.D. Zakharov, 30, 137–144 Zonneveld, J.P., T.W. Beatty, K.H. Williford, M.J. Orchard, and C.R. 2296 2291 Me´moires de Ge´ologie. McRoberts. 2010. Stratigraphy and Sedimentology of the lower 2297 2292 Zonneveld, J.P., and M.J. Orchard. 2002. Stratal relationships of the Black Bear Ridge section, British Columbia: candidate for the 2298 2293 Upper Triassic Baldonnel Formation, Williston Lake, northeast- base-Norian GSSP. Stratigraphy 7: 61–82. 2299 2294 ern British Columbia. Geological Survey of Canada Current 2295 Research 2002-A8: 1–13. Author Proof PROOF

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