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Clonal Colony in the Early Devonian Cnidarian Sphenothallus from Brazil

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Clonal Colony in the Early Devonian Cnidarian Sphenothallus from Brazil Clonal colony in the Early Devonian cnidarian Sphenothallus from Brazil HEYO VAN ITEN, JULIANA DE MORAES LEME, MARCELLO G. SIMÕES, and MARIO COURNOYER Van Iten, H., Leme, J.M., Simões, M.G., and Cournoyer, M. 2019. Clonal colony in the Early Devonian cnidarian Sphenothallus from Brazil. Acta Palaeontologica Polonica 64 (2): 409–416. The fossil record of polypoid cnidarians includes a number of taxa that were incorrectly identified as either tubiculous worms or plants. The holotype of the putative alga Euzebiola clarkei (Ponta Grossa Formation, Lower Devonian, Brazil), originally described under the name Serpulites sica, is re-described and re-figured as a species of Sphenothallus, a medu- sozoan cnidarian. Unlike Sphenothallus from other localities, the black, organic-walled Ponta Grossa specimen consists of a single parent tube that is confluent with the apical ends of at least 18 daughter tubes. The pattern of arrangement of the daughter tubes, which are arrayed in single file along the exposed face and the two thickened margins of the parent tube, partly resembles the whorl-like pattern of arrangement of colonial polyps of certain scyphozoan cnidarians. For these reasons, the Ponta Grossa Formation material figures prominently in the argument that Sphenothallus was a me- dusozoan cnidarian capable (in at least one species) of clonal budding. Key words: Cnidaria, Medusozoa, Scyphozoa, Hydrozoa, clonal budding, Devonian, Brazil. Heyo Van Iten [[email protected]], Department of Geology, Hanover College, Hanover, IN 47243, USA and Research Associate, Cincinnati Museum Center, Department of Invertebrate Paleontology, 1301 Western Avenue, Cincinnati, OH 45203, USA. Juliana de Moraes Leme [[email protected]], Department of Sedimentary and Environmental Geology, University of São Paulo, 05508-080, SP, Brazil. Marcello G. Simões [[email protected]], Department of Zoology, São Paulo State University, Botucatu campus, 18618-689, Botucatu, SP, Brazil. Mario Cournoyer [[email protected]], Musée de Paléontologie et de l’Évolution, 541 Congrégation Street, Montréal, Québec, Canada H3K 2J1. Received 28 November 2018, accepted 4 February 2019, available online 3 April 2019. Copyright © 2019 H. Van Iten et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License (for details please see http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. al. 2018), which hosts part of the record of the Cambrian Introduction Explosion (Yang et al. 2014). Together with the conula- riid Paraconularia sp. from the latest Ediacaran Tamengo Previous reports extending the known stratigraphical Formation of Brazil (Van Iten et al. 2014, 2016a; Parry et al. and paleogeographical ranges of the Paleozoic cnidarian 2017), Sphenothallus from Cambrian Stage 3 are among the Sphenothallus Hall, 1847 have increased the paleobiological oldest, polypoid medusozoan cnidarians in the fossil record significance of this extinct genus substantially (see Chang (Van Iten et al. 2014; Landing et al. 2018). et al. 2018; Landing et al. 2018 and references cited therein). Van Iten et al. (1992) first documented a suite of anatom- In addition to occurring in both oxic nearshore and dys- ical similarities uniquely shared by Sphenothallus, conulari- oxic offshore facies (e.g., Van Iten et al. 1996; Chang et al. ids and extant medusozoans, noting as well that similarities 2018), in such widely separated Cambrian and Ordovician shared by Sphenothallus and various non-cnidarian groups terranes as Laurentia (e.g., Van Iten et al. 1996, 2002), South are also shared with conulariids and extant medusozoans. China (e.g., Zhu et al. 2000; Li et al. 2004; Peng et al. 2005; The similarities thought to support assignment of both Van Iten et al. 2013; Muscente and Xiao 2015; Chang et Sphenothallus and conulariids to the subphylum Medusozoa al. 2018) and South Polar Gondwana (Fatka et al. 2012; Peterson, 1979 include: (i) soft body fully covered by a Fatka and Kraft 2013; Van Iten et al. 2016b), Sphenothallus tubular or steeply pyramidal, very finely lamellar, organic ranges downward into Cambrian Stage 3 (e.g., Chang et or organo-phosphatic periderm; (ii) periderm terminates Acta Palaeontol. Pol. 64 (2): 409–416, 2019 https://doi.org/10.4202/app.00576.2018 410 ACTA PALAEONTOLOGICA POLONICA 64 (2), 2019 54o 50o and extant medusozoans but also other invertebrates such O O O N A 70 60 50 O B 40 as graptolites and pterobranchs, is the capacity for asexual O o MT Brasília 10 16 budding. In support of their argument that at least some Sphenothallus formed clonal colonies, Van Iten et al. (1992: O 0 GO figs. 3, 4) presented two pieces of evidence: (i) a branching BRAZIL 20o MG specimen of Sphenothallus sp. from the Mississippian Bear 10O MS Brasília SP Paraná Gulch Formation (Montana, USA); and (ii) Clarke’s (1913: pl. Basin O 26: 16–18) shaded line drawings of a branching specimen of 20 24o Chaco Rio de PR Serpulites sica Salter, 1856 from the Lower Devonian Ponta Paraná Janeiro PARAGUAY Basin Curitiba Grossa Formation of southern Paraná State, Brazil. Clarke’s B 30O C Buenos Montevideo (1913) drawings depict what he interpreted as a single parent Aires SC 28o tube bearing at least 16 daughter tubes arrayed along the two 40O RS Porto Alegre ARGENTINA Atlantic margins of the parent, with the apical end of the daughter tubes Ocean 50O apparently lacking the basal holdfast and, instead, merging URUGUAY 0 200 km smoothly into the parent tube. Clarke (1913: 319) classified O 60 p-Cre arboniferous this specimen as a “problematum.” Later, Sommer (1954) units reexamined Clarke’s fossil, referring it to the new algal genus C 50O 49O and species Euzebiola clarkei. Interestingly, prior to 1913 the same specimen had been inspected by a paleobotanist, O N SP 24 PR who declared that is was “positively not a plant but possibly Arapoti SP-258 a hydroid (Clarke 1913: 319).” Moreover, Sommer (1954: 178) characterized his assignment of Clarke’s (1913) specimen to JJaguariaívaa the plant kingdom as merely a “taxonomic adventure”. Most recently, both this and the Bear Gulch Formation specimen were discussed by Landing et al. (2018) in their review paper Tibagi on the early evolution of coloniality in eumetazoans. These PR-340 Post-Devonianu nits authors added a third specimen, originally assigned to the PR-151 Ponta Grossa Formation genus Torellella Linnarsson, 1871 (Popov et al. 1989), to the Furnas Formation list of known or probable specimens of Sphenothallus show- Basin limit ing evidence of coloniality. Importantly, Van Iten et al. (1992) stated that the current O fault 25 Ponta whereabouts of Clarke’s (1913) branching specimen were Grossa road unknown. We here re-describe and re-illustrate Clarke’s Jaguariaíva-Arapoti railroad (1913) material, highlighting places on the branching spec- imen where diagnostic features of the genus Sphenothallus Curitiba and evidence of clonal budding are best preserved. We also BR-227 argue that both the branching specimen and similar, un- 0 10 20 km branched tubes from the Ponta Grossa Formation belong in the genus Sphenothallus. Fig. 1. A. Geographic location of the relevant geological sections within Paraná State, Brazil. B. Map showing the outcrop belt of the Lower Devonian Ponta Grossa Formation on the eastern margin of the Paraná Basin (from Simões et al. 2009). C. Geological map of the study area with Material and methods Jaguariaíva asterisked. Previously illustrated fossil specimens assigned by Clarke aborally in a sub-conical expansion that (in some cases) is (1913) to Serpulites sica occur on two cm-long slabs of mas- closed and floored by a very thin basal membrane; (iii) peri- sive siltstone that are housed in the paleontological collection derm exhibits (rarely) a transverse internal wall (schott) that of the National Mineral Agency (formerly the Department extends adorally along the inner surface of the periderm; of Mineral Production, DNPM) in Rio de Janeiro, south- and (iv) soft body includes two or more oral tentacles. Most eastern Brazil, under catalogue numbers DNPM 329 (the recently, Dzik et al. (2017) documented the presence in S. branching specimen; Clarke 1913: pl. 26: 16–18; Sommer ruedemanni (Lower Ordovician, China) of eight radially 1954: pl. 15: 2) and DNPM 1 (five partial unbranched tubes; arranged, seriated internal carinae similar to those present Clarke 1913: pl. 26: 15). Specific locality information is not in the periderm of certain coronate polyps but not known to provided in Clarke’s (1913) monograph and does not ac- occur in any other taxa except several species of conulariids company the reposited specimens. Thus, locating precisely (e.g., Van Iten et al. 1996). the sites that yielded these specimens is almost impossible. Another similarity, shared not only by Sphenothallus Nevertheless, information available in Clarke (1913: 62–67) VAN ITEN ET AL.—DEVONIAN SPHENOTHALLUS CNIDARIAN FROM BRAZIL 411 Epoch/Age Sequences/Lithostratigraphy Era Period Famennian PARAGUAY BRAZIL Late Frasnian Givetian Middle 634 Eifelian Devonian Emsian 458 Ponta Grossa Formation Early Pragian São Domingos Member Tibagi Member Lochkovian Paleozoic Jaguariaíva Member Pridoli Ludlow 337 Furnas Formation 323 ??? Wenlock Upper Middle/Lower Silurian conglomerate Landovery thickness (m) thickness (m) Fig. 2. Stratigraphic chart of the Silurian–Devonian interval in the Paraná Basin (modified from Assine et al. 1994; Sedorko et al. 2017). The exact level in the Jaguariaíva Member at which Clarke’s (1913) specimens of Sphenothallus were found is not known. indicates that his material comes from the Ponta Grossa contact (= transitional beds of Petri 1948) consisting of Formation (Lower Devonian) near the city of Ponta Grossa fine-grained sandstone with hummocky cross-stratification (Ponta Grossa County, Paraná State, southern Brazil). We and wavy cross-lamination that was originally deposited examined Clarke’s (1913) material under reflected light, in lower shoreface settings (Assine et al.
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