682
Journal of Paleontology 91(4):672–684
Chains, Zamora (2010); the rhombiferan Vizcainoia Zamora and Smith, 2012 from the Coulouma Formation [global Cambrian Series 3, lower Drumian], Montagne Noire [France], Zamora and Smith (2012)). The Jbel Warwmast Formation, the Murero Formation, and the Coulouma Formation consist of upper offshore fine siliciclastic sediments (siltstone to clay- stone) deposited under low or moderately low energy conditions in a temperate mixed platform (Liñan and Mergl, 2001; Landing et al., 2006; Álvaro et al., 2008). The closely related and con- temporaneous lepidocystoids occur in obrution deposits in the Impure Carbonate Facies in the Emigsville Member of the Kinzers Formation (global Cambrian Series 2, Stage 4) in the Conestoga Valley (southern Pennsylvania), interpreted as an offshore environment with active bottom currents in a mixed platform (Skinner, 2005). An undetermined lepidocystoid was mentioned in the transgressive claystone-dominated Issafen Formation [global Cambrian Series 2, Stage 4] in the Central Anti-Atlas (Morocco) (Smith et al., 2013). The (par)auto- chthonous occurrences of the felbabkacystids and lepidocystoids in the deeper facies in the Paradoxides (P.) paradoxissimus gracilis Zone [global Cambrian Series 3, mid-Drumian] suggest that the morphological innovation of the overgrown vaulted oral area might have occurred in an offshore and quiet environment. All occurrences of lepidocystoids, felbabkacystids, and Medi- terranean eocrinids-lichenoids are related to relatively deep (up to mid offshore) lithofacies in transgressive and/or drowning plat- form contexts (Skinner, 2005; Landing et al., 2006; Fatka and Szabad, 2014). The echinoderm plasticity and geographic dis- tribution, even at low taxonomic levels, might be controlled by environmental changes such as the water energy, the water depth, and the substrate consistency (Guensburg and Sprinkle, 1992; Sprinkle and Guensburg, 1995; Álvaro et al., 2013; Smith et al., 2013). The morphological innovation and the high diversity in offshore environments during the early mid-Cambrian times seem to dispute the onshore-offshore pattern for the early echinoderm diversification (Jablonski et al., 1983; Sepkoski, 1991; Smith et al., 2013).
Acknowledgments
All authors contributed to this study. This study was supported by MSM 0021620855 and the Czech Science Foundation through the PRVOUK P44 of the Ministry of Education, Youth and Sports of Czech Republic. It is a contribution to the group “Lithosphère-Océan-Atmosphère” of the UMR CNRS5563/ IRD234/UPS/CNES GET (EN), to the SYNTHESYS projects SE-TAF-2924 (EN) and CZ-TAF-6049 (BL), and to the UMR CNRS 5276 LGLTPE (BL). The authors are particularly grate- ful to T. E. Guensburg and R. L. Parsley for their very helpful comments, and to S. Zamora, C. D. Sumrall, and J. Sprinkle for their constructive remarks, which greatly helped to improve the manuscript.
Accessibility of supplemental data
Data available from the Dryad Digital Repository:
http://dx.doi. org/10.5061/
dryad.sg931
References
Álvaro, J.J., Bauluz, B., Subias, I., Pierre, C., and Vizcaïno, D., 2008, Carbon chemostratigraphy of the Cambrian-Ordovician transition in a midlatitude mixed platform, Montagne Noire, France: Geological Society of America Bulletin, v. 120, p. 962–975.
Álvaro, J.J., Zamora, S., Clausen, S., Vizcaïno, D., and Smith, A.B., 2013, The role of abiotic factors in the Cambrian Substrate Revolution: A review from the benthic community replacements of West Gondwana: Earth-Science Reviews, v. 118, p. 69–82.
Ausich,W.I., 2001, Echinodermtaphonomy, in Jangoux, M., and Lawrence, J.M., eds., Echinoderm Studies, v. 6. Lisse, A.A. Balkema, p. 171–227.
Babcock, L.E., Rees, M.N., Robison, R.A., Langenburg, E.S., and Peng, S., 2004, Potential Global Standard Stratotype-section and Point (GSSP) for a Cambrian stage boundary defined by the first appearance of the trilobite Ptychagnostus atavus, Drum Mountains, Utah, USA: Geobios, v. 37, p. 149–158.
Barrande, J., 1846, Notice préliminaire sur le Système silurien et les trilobites de Bohème: Leipzig, Hirschfeld, 97 p.
Barrande, J., 1887, Systême Silurien du centre de la Bohême. Part I: Recherches paléontologiques, v. 7, Classes des Echinodermes, sec. 1, Ordre des Cystidées: Leipzig, Rivnac, Prague/Gerhard, 396 p.
Bottjer, D.J., and Ausich, W.I., 1986, Phanerozoic development of tiering in soft substrate suspension-feeding communities: Paleobiology, v. 12, p. 400–420.
Bottjer, D.J., Hagadorn, J.W., and Dornbos, S.Q., 2000, The Cambrian substrate revolution: GSA Today, v. 10, p. 1–7.
Branson, E.B., and Peck, R.E., 1940, A new cystoid from the Ordovician of Oklahoma: Journal of Paleontology, v. 14, p. 89–92.
Brett, C.E., Moffat, H.A., and Taylor, W., 1997, Echinoderm taphonomy, taphofacies, and Lagerstätten, in Waters, J.A., and Maples, C.G., eds., Geobiology of Echinoderms: Paleontological Society Papers, v. 3, p. 147–190.
Broadhead, T.W., 1982, Reappraisal of class Eocrinoidea (Echinodermata), in Laurence, J.M., ed., Echinoderms: Proceedings of the 4th International Echinoderm Conference; Rotterdam, A.A. Balkema, p. 125–131.
Callaway, C., 1877, On a new area of Upper Cambrian rocks in South Shrop- shire, with a description of new fauna: Quarterly Journal of the Geological Society of London, v. 33, p. 652–672.
Chauvel, J., 1941, Recherches sur les Cystoïdes et les Carpoïdes armoricains: Mémoires de la Société Géologique et Minéralogique de Bretagne, v. 5, p. 1–286.
David, B., Lefebvre, B., Mooi, R., and Parsley, R.L., 2000, Are Homalozoans echinoderms? An answer from the extraxial-axial theory: Paleobiology, v. 26, p. 529–555.
Dornbos, S.Q., 2006, Evolutionary palaeoecology of early epifaunal echino- derms: Response to increasing bioturbation levels during the Cambrian radiation: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 237, p. 225–239.
Durham, J.W., 1968, Lepidocystoids, in Moore, R.C., ed., Treatise on Invertebrate Paleontology, Part S, Echinodermata 1(2): New York and Lawrence: Geological Society of America and the University of Kansas, p. S631–S634.
Eichwald, C.L. von, 1840, Über das silurische Schichtensystem in Esthland: St. Petersburg: Medizinische Akademie, 222 p.
Fatka, O., 2006, Biostratigraphy of the Jince Formation (middle Cambrian) in the Příbram-Jince Basin: Historical review: Acta Universitatis Carolinae - Geologica, v. 47, p. 53–61.
Fatka, O., and Kordule, V., 1985, Etoctenocystis bohemica gen. et sp. nov., new Ctenocystoid from Czechoslovakia (Echinodermata, middle Cambrian): Věstník Ústředního ústav geologického, v. 60, p. 225–231.
Fatka, O., and Kordule, V., 1990, Vyscystis ubaghsi gen. et sp. nov., imbricate eocrinoid from Czechoslovakia (Echinodermata, middle Cambrian): Věstník Ústředního ústav geologického, v. 65, p. 315–323.
Fatka, O., and Kordule, V., 1991, Akadocrinus knizki sp. nov., gogiid eocrinoids from Czechoslovakia (Echinodermata, middle Cambrien): Věstník Ústředního ústav geologického, v. 66, no. 4, p. 239–246.
Fatka, O., and Kordule, V., 2001, Asturicystis havliceki sp. nov. (Echinodermata, Homostelea) form the middle Cambrian of Bohemia (Barrandian area, Czech Republic): Journal of Czech Geological Survey, v. 46, p. 189–194.
Fatka, O., and Mergl, M., 2009, The “microcontinent” Perunica: Status and story 15 years after conception: Geological Society, London, Special Publica- tions, v. 325, p. 65–101.
Fatka, O., and Szabad, M., 2014, Cambrian biostratigraphy in the Příbram-Jince Basin (Barrandian area, Czech Republic): Bulletin of Geosciences, v. 89, p. 413–429.
Page 1 |
Page 2 |
Page 3 |
Page 4 |
Page 5 |
Page 6 |
Page 7 |
Page 8 |
Page 9 |
Page 10 |
Page 11 |
Page 12 |
Page 13 |
Page 14 |
Page 15 |
Page 16 |
Page 17 |
Page 18 |
Page 19 |
Page 20 |
Page 21 |
Page 22 |
Page 23 |
Page 24 |
Page 25 |
Page 26 |
Page 27 |
Page 28 |
Page 29 |
Page 30 |
Page 31 |
Page 32 |
Page 33 |
Page 34 |
Page 35 |
Page 36 |
Page 37 |
Page 38 |
Page 39 |
Page 40 |
Page 41 |
Page 42 |
Page 43 |
Page 44 |
Page 45 |
Page 46 |
Page 47 |
Page 48 |
Page 49 |
Page 50 |
Page 51 |
Page 52 |
Page 53 |
Page 54 |
Page 55 |
Page 56 |
Page 57 |
Page 58 |
Page 59 |
Page 60 |
Page 61 |
Page 62 |
Page 63 |
Page 64 |
Page 65 |
Page 66 |
Page 67 |
Page 68 |
Page 69 |
Page 70 |
Page 71 |
Page 72 |
Page 73 |
Page 74 |
Page 75 |
Page 76 |
Page 77 |
Page 78 |
Page 79 |
Page 80 |
Page 81 |
Page 82 |
Page 83 |
Page 84 |
Page 85 |
Page 86 |
Page 87 |
Page 88 |
Page 89 |
Page 90 |
Page 91 |
Page 92 |
Page 93 |
Page 94 |
Page 95 |
Page 96 |
Page 97 |
Page 98 |
Page 99 |
Page 100 |
Page 101 |
Page 102 |
Page 103 |
Page 104 |
Page 105 |
Page 106 |
Page 107 |
Page 108 |
Page 109 |
Page 110 |
Page 111 |
Page 112 |
Page 113 |
Page 114 |
Page 115 |
Page 116 |
Page 117 |
Page 118 |
Page 119 |
Page 120 |
Page 121 |
Page 122 |
Page 123 |
Page 124 |
Page 125 |
Page 126 |
Page 127 |
Page 128 |
Page 129 |
Page 130 |
Page 131 |
Page 132 |
Page 133 |
Page 134 |
Page 135 |
Page 136 |
Page 137 |
Page 138 |
Page 139 |
Page 140 |
Page 141 |
Page 142 |
Page 143 |
Page 144 |
Page 145 |
Page 146 |
Page 147 |
Page 148 |
Page 149 |
Page 150 |
Page 151 |
Page 152 |
Page 153 |
Page 154 |
Page 155 |
Page 156 |
Page 157 |
Page 158 |
Page 159 |
Page 160 |
Page 161 |
Page 162 |
Page 163 |
Page 164 |
Page 165 |
Page 166 |
Page 167 |
Page 168 |
Page 169 |
Page 170 |
Page 171 |
Page 172 |
Page 173 |
Page 174 |
Page 175 |
Page 176 |
Page 177 |
Page 178 |
Page 179 |
Page 180 |
Page 181 |
Page 182 |
Page 183 |
Page 184 |
Page 185 |
Page 186 |
Page 187 |
Page 188 |
Page 189 |
Page 190 |
Page 191 |
Page 192 |
Page 193 |
Page 194 |
Page 195 |
Page 196 |
Page 197 |
Page 198 |
Page 199 |
Page 200 |
Page 201 |
Page 202 |
Page 203 |
Page 204 |
Page 205 |
Page 206 |
Page 207 |
Page 208 |
Page 209 |
Page 210 |
Page 211 |
Page 212 |
Page 213 |
Page 214 |
Page 215 |
Page 216 |
Page 217 |
Page 218 |
Page 219 |
Page 220 |
Page 221 |
Page 222 |
Page 223 |
Page 224 |
Page 225 |
Page 226 |
Page 227 |
Page 228 |
Page 229 |
Page 230 |
Page 231 |
Page 232 |
Page 233 |
Page 234 |
Page 235 |
Page 236 |
Page 237 |
Page 238 |
Page 239 |
Page 240 |
Page 241 |
Page 242 |
Page 243 |
Page 244 |
Page 245 |
Page 246 |
Page 247 |
Page 248 |
Page 249 |
Page 250 |
Page 251 |
Page 252 |
Page 253 |
Page 254 |
Page 255 |
Page 256 |
Page 257 |
Page 258 |
Page 259 |
Page 260 |
Page 261 |
Page 262 |
Page 263 |
Page 264 |
Page 265 |
Page 266 |
Page 267 |
Page 268 |
Page 269 |
Page 270 |
Page 271 |
Page 272 |
Page 273 |
Page 274 |
Page 275 |
Page 276 |
Page 277 |
Page 278 |
Page 279 |
Page 280 |
Page 281 |
Page 282 |
Page 283 |
Page 284 |
Page 285 |
Page 286 |
Page 287 |
Page 288
