Palaeogeography, Palaeoclimatology, Palaeoecology 460 (2016) 170–178 Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo Palaeobiogeography, palaeoecology and evolution of Lower Ordovician conulariids and Sphenothallus (Medusozoa, Cnidaria), with emphasis on the Fezouata Shale of southeastern Morocco Heyo Van Iten a,b,⁎, Lucy Muir c, Marcello G. Simões d, Juliana M. Leme e, Antonio C. Marques f,g, Naomi Yoder h a Department of Geology, Hanover College, Hanover, IN 47243, USA b Cincinnati Museum Center, Department of Invertebrate Paleontology, 1301 Western Avenue, Cincinnati, OH 45203, USA c Department of Geology, Amgueddfa Cymru — National Museum of Wales, Cathays Park, Cardiff CF10 3NP, Wales, UK d Department of Zoology, São Paulo State University, Rubião Junior District, 18618-000 Botucatu, São Paulo, Brazil e Geosciences Institute, University of São Paulo, R. Lago 562, 05508-080, São Paulo, Brazil f Biosciences Institute, University of São Paulo, R. Matão 101, 05508-090 São Paulo, Brazil g Center for Marine Biology, University of São Paulo, São Sebastião, Brazil h Department of Marine Science, University of Southern Mississippi, Stennis Space Center, MS 39529, USA article info abstract Article history: The fossil record of conulariids (Cnidaria, Scyphozoa) extends downward into the topmost part of the Ediacaran Received 31 August 2015 System, but the first appearance of diverse, widespread conulariids is in siliciclastic rock units of Early Ordovician Received in revised form 4 March 2016 age, which collectively host at least six conulariid genera. Some of these same units also contain Sphenothallus,a Accepted 10 March 2016 probable medusozoan that frequently co-occurs with conulariids in Ordovician and younger deposits. Lower Available online 15 March 2016 Ordovician conulariid localities are distributed among five (originally) Southern Hemisphere terranes, namely Core Gondwana (Archaeoconularia, Eoconularia and Teresconularia), Armorica (Conularia azaisi), Avalonia Keywords: Conulariids (Archaeoconularia, Eoconularia and Exoconularia), Perunica (Archaeoconularia, Conularia and Conulariella) and Sphenothallus South China (Conulariella). C. azaisi, currently known from the Southern Montagne Noire (France), probably rep- Medusozoa resents a new genus. Sphenothallus occurs in South China, North China (Korea), Armorica (Southern Montagne Lower Ordovician Noire) and Core Gondwana (Morocco). In southeastern Morocco, Burgess Shale-type Konservat-Lagerstätten in Gondwana the Fezouata Shale (Tremadocian–Floian) yield Archaeoconularia sp., Eoconularia sp. and at least one species of Fezouata Sphenothallus. This low-diversity conulariid assemblage is most similar to the Tremadocian assemblage of Wales (Avalonia), which likewise consists of a single species each of Archaeoconularia and Eoconularia. In the Fezouata Shale, Archaeoconularia sp. and Eoconularia sp. frequently occur in monospecific mass associations. Such associations probably represent an original clumped distribution on the seafloor. Additionally, some Eoconularia sp. occur in V-like pairs or radial clusters, and also some specimens were attached at the apical end to a phosphatic brachiopod or to a corner sulcus of a larger specimen of Eoconularia sp. Similar conulariid/ brachiopod associations, consisting of Conularia trentonensis and Onniella sp., occur in the Upper Ordovician (Katian) Collingwood Shale of southern Ontario, Canada. © 2016 Elsevier B.V. All rights reserved. 1. Introduction Cambrian systems, respectively (e.g. Zhu et al., 2000; Li et al., 2004; Van Iten et al., 2014a). Sphenothallus has been reported widely from Conulariids (order Conulariida Miller and Gurley, 1896)and the Cambrian System (see for example Fatka and Kraft, 2013; Sphenothallus Hall, 1847, two extinct groups of medusozoan cnidarians Muscente and Xiao, 2015), but thus far Cambrian conulariids are repre- that produced a finely lamellar, organo-phosphatic periderm (Van Iten sented solely by Baccaconularia Hughes et al., 2000 from the Furongian et al., 1992, 2006a), first appear in the uppermost Ediacaran and lower Saint Lawrence Formation (Upper Mississippi Valley, USA), though conulariids may occur in Cambrian Stage 1 (see Van Iten et al., 2010 and references cited therein), and Paraconularia sp. has been found in the latest Ediacaran Tamengo Formation of Brazil (Van Iten et al., ⁎ Corresponding author at: Department of Geology, Hanover College, Hanover, IN 2014a). The last occurrence is particularly intriguing as recent cladistic 47243, USA. analyses (Leme et al., 2008; Van Iten et al., 2014b)positthat E-mail addresses: [email protected] (H. Van Iten), [email protected] (L. Muir), [email protected] (M.G. Simões), [email protected] (J.M. Leme), Paraconularia Sinclair, 1940 was a relatively apical branch on the [email protected] (A.C. Marques), [email protected] (N. Yoder). conulariid tree. In a number of Ordovician and younger rock units, http://dx.doi.org/10.1016/j.palaeo.2016.03.008 0031-0182/© 2016 Elsevier B.V. All rights reserved. (e.g. well-preserved fossils ranging from micro-invertebratesbeen to sampled chordates intensively, yielding tens of thousands of exceptionally number UMMP); and Yale UniversityMichigan Peabody Museum Museum of (New Paleontology Haven, (Ann Arbor,(Cardiff, USA; UK); specimen Natural pre HistoryFrance Museum (sample (London, UK); numberUCBL-FSL); University pre of Department ofEarth Earth Sciences, Sciences, University Montpellier of University, Lyon, France (sample number pre 2016 the upper part and isTremadocian mid in Floian in age, age and ( (60 the m other thick) of occurs whichof conulariids (15 in collectively m the occur in thick) lowerRoy both occurs et of part in al., these of 2010; intervals, Martin one the etfrom of al., formation Burgess which in Shale-type and press biotas of isarthropods early late or and middle other Cambrian taxa, agetwo some ( exceptionally preserved of fossil intervals which (EPFs) yielding were soft-bodied originally described Group of northwestern Argentina ( discoveries of Lower Ordovician conulariidsby are palaeoenvironmental in the or SantaSphenothallus Victoria taphonomic artifacts. TheOverstreet most et al., recent 2003 fossiliferous dolostones depositedlow-latitude in terranes restricted such environmentsthe as fact (e.g. Laurentia that are the Lower dominatedpalaeolatitudes Ordovician by ( rock sparsely records ofexception of the North relatively China, few generally were situatedpredominantly at mid- Gondwanan to very and high are peri-Gondwanan, from which, (originally) withcurrently Southern the Hemisphere known terranes Lower and Ordovician localities, conulariids and Marrakesh, Morocco (sampleseven institutions: Department of Earth number Sciences, Cadi Ayyad University, pre on direct examination of fossil specimens in collections of the following 2. Material and methods attached to other conulariid specimens. attachment to phosphaticexceptionally brachiopods preserved conulariids and exhibiting evidence conulariidsaddition of to original apparently Burgess apical Shale-type organisms, the Fezouata Shale has yielded some of whichthroughout also much host of the world, is inpreferences. Lower Ordovician formations, and possibly being closely related to eachrare other or phylogenetically, conulariids absent (e.g. benthos such as corals, crinoids and rhynchonelliformin brachiopods are some cases inconulariids and relatively restricted facies in which normal marine Canada; is extremely rare in the mid Cambrian Burgess Shale (British Columbia, with ( Van Iten et al., 2013 the Tonggao Formation of Guizhou Province, South China ( rock unit in Africa knownattention to to contain conulariids the and Fezouata Shale,relationships which at among present conulariid is thein genera only the Ordovician context ( of aecology parsimony-based and cladistic evolution hypothesis of Lower of Ordovician phylogenetic conulariids and conulariid. other Cambrian Konservat-Lagerstätte has produced even a single Van Roy et al., 2015 The present study is based on review of the relevant literature and The Thepresentarticleaddresseskeyasp – Sphenothallus Caron and Jackson, 2008; Zhao et al., 2010 Sphenothallus in this issue fi rst appearance of multiple genera of conulariids, in localities Van Iten et al., 2002 in these regions may be, at least to some extent, caused Sphenothallus Cocks and Torsvik, 2004 ). Many Burgess Shale-type Cambrian faunas have . The Tremadocian ), and the Fezouata Shale of southeastern Morocco may have shared similar palaeoenvironmental Van Iten et al., 1996, 2012 ). In the last two formations, conulariids co-occur ), suggesting that the lack of conulariids and Sphenothallus occur in close association with each other, fi x UM); National Museum of Wales ), but to date neither this nor any Teresconularia – H. Van Iten et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 460 (2016) 170 ). Floian Fezouata Shale contains ects of the distribution, palaeo- Sphenothallus Gutiérrez Marco and Martin, ( Fig. 1 Table 1 ). In part this may re fi x AA); Department of ). Thus, in addition to ). We pay particular ; ). ). Interestingly, all Leme et al., 2003 Sphenothallus Sphenothallus and two species Sphenothallus Sphenothallus Conulariella fl Van .In ect sp. fi fi ), x x ; , Table 1 Distribution of Lower Ordovician conulariids and Sphenothallus [in brackets]. An asterisk (*) next to a species name indicates that the occurrence