Journal of Paleontology, 91(4), 2017, p. 735–754 Copyright © 2017, The Paleontological Society 0022-3360/17/0088-0906 doi: 10.1017/jpa.2017.7


The first Ordovician cyclocystoid (Echinodermata) from Gondwana and its morphology, paleoecology, taphonomy, and paleogeography


Mike Reich,1,2,3 James Sprinkle,4 Bertrand Lefebvre,5 Gertrud E. Rössner,1,2,3 and Samuel Zamora6


1SNSB – Bavarian State Collection of Palaeontology and Geology, Richard-Wagner-Strasse 10, 80333 München, Germany ⟨m.reich@lrz.uni-muenchen.de⟩, ⟨g.roessner@lrz.uni-muenchen.de⟩ 2Department of Earth and Environmental Sciences, Division of Palaeontology and Geobiology, Ludwig-Maximilians-Universität München,


Richard-Wagner-Strasse 10, 80333 München, Germany ⟨mike.reich@lmu.de⟩ 3GeoBio-Center, Ludwig-Maximilians-Universität München, Germany 4Department of Geological Sciences, Jackson School of Geosciences, University of Texas, 1 University Station C1100, Austin, Texas


78712-0254, USA ⟨echino@jsg.utexas.edu⟩ 5UMR CNRS 5276 LGLTPE, Université Lyon 1, 2 rue Raphaël Dubois, 69622 Villeurbanne cedex, France ⟨bertrand.lefebvre@univ-lyon1.fr⟩ 6Instituto Geológico y Minero de Espana, C/Manuel Lasala 44, 9 B, 50006 Zaragoza, Spain ⟨samuel@unizar.es⟩


Abstract.—Moroccodiscus smithi represents a new cyclocystoid genus and species based on moldic specimens from the Middle Ordovician Taddrist Formation (Darriwilian) of SE Morocco. This represents the earliest articulated mem- ber of the Cyclocystoidea and is the first complete cyclocystoid described from the Ordovician of Gondwana, as well as the first cyclocystoid ever recorded from Africa. The anatomy and morphology of this new species were studied using a combination of conventional paleontological methods and nondestructive X-ray computed tomography. Because Moroccodiscus differs from other cyclocystoids, in particular by lacking cupules attached to the marginal ossicles, it is assigned to the new family Moroccodiscidae. This new taxon illustrates the relatively poorly known early diversification of these enigmatic extinct echinoderms and sheds light on the mode of life of cyclocystoids, including injuries to plate circlets during early ontogeny and folding of these disk-like specimens at the time of death. The overall thecal shape was very similar in cyclocystoids and many domal edrioasteroids, probably because they were both sessile or attached, benthic, suspension feeders. However, many oral surface, ambulacral, and marginal ring features had become very different, indicating that these two groups had either converged because of similar life modes or were only distantly related sister groups.


Introduction


Cyclocystoids are a relatively small extinct class of circular, flattened echinoderms that have been known for more than 160 years. Twenty genera and about 40 valid species have been named (Reich and Kutscher, 2010; Sprinkle et al., 2015), from articulated and disarticulated material, ranging from the Middle Ordovician to the early Carboniferous (Heaslip, 1969; Smith and Paul, 1982; Ressmeyer and Frest, 1983; Berg-Madsen, 1987; Fluegeman and Orr, 1990; Haude and Thomas, 1994; Boczarowski, 2001; Sevastopulo, 2002). Most specimens come from Laurentia and Baltica (Lefebvre et al., 2013), but a few are known from Avalonia and Laurussia. Jell and Jell (1999) pub- lished the only report of a cyclocystoid plate from Gondwana (Upper Devonian of Australia). However, our new cyclocystoid material described here represents the second report (and first articulated Ordovician material) of this echinoderm group from Gondwana. Cyclocystoid morphology has been controversial because


well-preserved specimens that show all parts of the animal are rare (Paul, 1982), and several different interpretations of their morpho- logy and life mode have been proposed (Salter and Billings, 1858;


Foerste, 1920; Sieverts-Doreck, 1951; Kesling, 1963, 1966). In addition, despite the abundance of cyclocystoids in Ordovician strata (Table 1) in comparison to the Silurian– Carboniferous record, our understanding of their origin and phylogeny lags well behind that of many other echinoderm groups. Smith and Paul (1982) produced a comprehensive monograph of the class using a large number of well-preserved museum specimens and established six new genera. We agree with many of their conclusions involving systematics, bio- stratigraphy, morphology, diversity, phylogeny, and perhaps biogeography, but disagree with their interpretation of orienta- tion and feeding mode. The recent discovery of a new cyclocystoid (Moroccodiscus


smithi) fromthe Middle Ordovician ofMorocco shows somewhat different morphology and additional evidence about the likely living orientation of cyclocystoids. Here we describe the eight specimens of this new taxon that have been assembled by the authors, showing different morphology in comparison to all other known cyclocystoid species, genera, and families. Moroccodiscus smithi from the Middle Ordovician of Morocco represents the oldest articulated member of the Cyclocystoidea as well as the first record of this group from Africa.


735


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