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An Upper Mississippian Echinoderm Microfauna from the Genicera Formation of Northern León (Carboniferous, Cantabrian Mountains, N Spain)

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An Upper Mississippian Echinoderm Microfauna from the Genicera Formation of Northern León (Carboniferous, Cantabrian Mountains, N Spain) SPANISH JOURNAL OF PALAEONTOLOGY An Upper Mississippian echinoderm microfauna from the Genicera Formation of northern León (Carboniferous, Cantabrian Mountains, N Spain) Joachim PABST* & Hans-Georg HERBIG Institut für Geologie und Mineralogie, Universität zu Köln. Zülpicher Str. 49a, D-50674 Köln. [email protected]; [email protected] * Corresponding author Pabst, J. & Herbig, H. 2020. An Upper Mississippian echinoderm microfauna from the Genicera Formation of northern León (Carboniferous, Cantabrian Mountains, N Spain). [Microfauna de equinodermos del Misisipiense Superior de la Formación Genicera del norte de León (Carbonífero, Cordillera Cantábrica, N de España)]. Spanish Journal of Palaeontology, 35 (1), 47-76. Manuscript received 30 January 2019 https://doi.org/10.7203/sjp.35.1.17116 Manuscript accepted 12 November 2019 © Sociedad Española de Paleontología ISSN 2255-0550 ABSTRACT RESUMEN For the fi rst time an echinoderm microfauna is recorded from Por primera vez se describe una microfauna de equinodermos the cephalopod limestone facies (‘griotte facies’) of the lower de las calizas nodulosas con cefalópodos (“griotte facies”) Carboniferous (Mississippian) Genicera Fm. (Alba Fm.). de la Formación Genicera (Formación Alba) del Carbonífero The formation is widespread in the Cantabrian Mountains inferior (Misisipiense). Si bien esta formación es de in NW Spain, but the ossicles are from some sections in considerable extensión en la Cordillera Cantábrica en el the surroundings of the Bernesga valley in northern León. norte de España, los oscículos descritos solo proceden de They have been derived from insoluble acetic acid residues unas secciones de los alrededores del valle de Bernesga al from samples of the upper and especially of the uppermost norte de León. Tras un tratamiento con ácido acético, éstos part of the formation (Canalón Mb. and Millaró Beds). The han sido identifi cados en los residuos insolubles de pruebas microfauna include taxonomically treated wheel-shaped de la parte superior y especialmente de los últimos niveles ossicles, sieve-plates and rods of holothurians, goniodonts of de dicha formación (Miembro Canalón y Capas de Millaró). ophiocistioids, and ophiuroid and stenuroid skeletal elements. La microfauna incluye oscículos con forma de rueda, placas From the Palaeozoic of Spain Ophiocistioidea, Stenuroidea, perforadas y bastones de los holoturoideos, goniodontos de Apodida (Holothuroidea), and allagecrinids (microcrinoids) los ofi ocistioideos y elementos de ofi uroideos y stenuroideos, are first reported. Here, we describe two new species: todos ellos taxonomicamente estudiados. Varios elementos de Linguaserra heidii n. sp. (Ophiocistioidea) and Calclyra la microfauna representan hallazgos inéditos del Paleozoico bifida n. sp. (Ophiuroidea). Indeterminate echinoderm de España, incluso de sus correspondientes rangos más altos remains are also fi gured and discussed. The fi ndings stress como Ofi ocistioidea, Stenuroidea, Apodida (Holothuroidea), y the importance of a diverse but still poorly documented allagecrinidos (microcrinoideos). En este trabajo describimos 48 PABST & HERBIG echinoderm fauna in Upper Mississippian psychrospheric dos nuevas especies: Linguaserra heidii n. sp. (Ophiocistioidea) pelagic environments. y Calclyra bifi da n. sp. (Ophiuroidea). Además se expone y discute unos restos de equinodermos indeterminados. Estos Keywords: Viséan, Serpukhovian, cephalopod limestone, hallazgos reivindican la importancia de una diversa fauna carbonate microfacies, echinoderm ossicles, taxonomy. de equinodermos todavía insufi cientemente documentada del ámbito psicrosférico pelágico del Misisipiense Superior. Palabras clave: Viséense, Serpukhoviense, calizas con cefalópodos, microfacies de carbonatos, oscículos de equinodermos, taxonomía. 1. INTRODUCTION limestone”, resp. “marbre griotte”. Pelagic dwelling organisms and taxa preferring deeper-water environments The Cantabrian Mountains form the most external zone predominate. These are goniatites, cyathaxonid corals, of the northern branch of the Spanish Variscides, named trilobites characteristic of the Kulm facies, and conodonts; Cantabrian Zone (Lotze, 1945), which is characterized by opposed brachiopods are scarce (see review in Sánchez a complex pattern of thrust slices and nappes (Julivert, de Posada et al., 1996). Later contributions added to the 1971). During most of the Mississippian period that knowledge of bivalves (Amler & Winkler Prins, 1999) foreland basin passed through the pre-orogenic starved and agglutinating foraminifers (Balthasar & Amler, 2003; basin phase characterized by pelagic deeper water deposits Herbig et al., 2011). (Kullmann et al., 1977; Marcos & Pulgar, 1982; Colmenero Macroscopic remains of echinoderms are exceedingly et al., 2002). During the mid and late Tournaisian, some rare within the formation, although articulated pelmatozoan meters thick black shales developed (Vegamián Fm.) stem ossicles occur in places, and disarticulated remains (Comte, 1959). They are overlain by condensed pelagic are quite common in thin sections. Herbig (1982, 1994) nodular cephalopod limestone (‘griotte’) of the Genicera described an at least in part autochthonous monospecifi c Fm. (Wagner et al., 1971), respectively Alba Fm. (Comte, crinoid fauna with Balearocrinus cantabricus Herbig, 1959), including locally a subordinate package of siliceous 1982, a genus, which is also known from time-equivalent shales and cherts. The formation does not exceed a deepwater deposits of Menorca (Bourrouilh & Termier, thickness of 55 m in spite of its extended stratigraphic 1973) and the Bechar Basin of Algeria (Webster et al., range, reaching from the diachronous base between latest 2004). A further crinoid cup from the Genicera Fm. Tournaisian and earliest Viséan (Scaliognathus anchoralis of northern León was described by Breimer (1962, to Gnathodus texanus zone) to the late Serpukhovian not figured); Rhabdocrinus scotocarbonarius from (Gn. truyolsi zone). Among numerous studies, some contemporaneous cephalopod limestones of the Palentine relevant biostratigraphic papers are from van Adrichem realm (Morris, 1992) belongs to Balearocrinus cantabricus Boogaert (1967), Higgins (1971), Higgins & Wagner- (Herbig, 1994). Gentis (1982), García-López & Sanz-López (2002) and Zamora (2016) reviewed the Palaeozoic echinoderms Nemyrovska (2005). In the last decade, uppermost Viséan from northern Spain and listed the few isolated and Serpukhovian conodonts and smaller foraminifers of Carboniferous fi nds. Except from crinoids, descriptions the Genicera Fm. gained attention due to their importance of Pennsylvanian blastoids (Sieverts-Doreck, 1951; for definition of a GSSP at the Viséan–Serpukhovian Joysey & Breimer, 1963), asterozoans and echinoids boundary (e.g. Sanz-López et al., 2007; Cózar et al., 2016; (Bless & Sánchez de Posada, 1971) exist. Bless & Sanz-López et al., 2019). The Genicera Fm. and its lateral Sánchez de Posada (1971) also mentioned holothurian equivalents, e.g. the Carrión Fm. of Nemyrovska (2005) ossicles without further specifi cation. First short accounts from the Palentine realm, are widespread in all tectonic on the present fauna were published by Pabst & Herbig units of the Cantabrian Mountains containing Mississippian (2018, 2019). Herein, we document these fi rst mainly rocks. Its sediments represent the largest preserved late non-crinoid echinoderm remains from the Genicera Fm., Palaeozoic pelagic carbonate platform of the European– namely disarticulated holothurian, ophiuroid, ophiocistioid North African Variscides. However, analogous facies and stenuroid ossicles as well as other indeterminate are also widespread in other South Armorican Massifs echinoderm ossicles in detail. It has to be stressed that (Pyrenees, Catalonian Coastal Ranges, Menorca). these are the first reports of ophiocistioids, of apodid The limestone of the Genicera Fm. yielded a wealth holothurians, of some “arcaic ophiuroids” (Furcaster of fossils, first mentioned by Schulz (1837), Prado & among Oegophiuroidea, Stenuroidea) and of microcrinoids de Verneuil (1850) and Barrois (1882) from the “griotte (allagecrinids) from the Palaeozoic of Spain. Apart AN UPPER MISSISSIPPIAN ECHINODERM MICROFAUNA FROM THE GENICERA FORMATION OF NORTHERN LEÓN ... 49 from two Upper Devonian occurrences of actinopodian ammonoid and trilobite fi ndings of Higgins et al. (1964) holothurians (?Microantyx Kornicker & Imbrie, 1958 and and Gandl (1977). The section north of Rabanal de Fenar, one eucaudinid) described by García-López & Truyols southeastern Alba Syncline, hitherto remained unstudied. (1974), no taxonomical study on holothuroid ossicles from A near-by section is noteworthy due to an important, local the Palaeozoic of Spain exists at all. debrisfl ow deposit at the top of the Genicera Fm. reworking All ossicles have been derived from three sections in limestone clasts of the San Adrián Mb. (Keller et al., 2008). northern León during an ongoing study linking carbonate In the type locality of the Genicera Fm. Wagner et al. microfacies and agglutinating foraminifers. (1971) distinguished three members, from below Gorgera Mb. (cephalopod limestone), Lavandera Mb. (shales and cherts) and Canalón Mb. (cephalopod limestone). However, this subdivision varies considerably in the Asturo-Leonese 2. GEOLOGICAL AND STRATIGRAPHICAL facies realm, as exemplified by sections in the wider surroundings of the Bernesga Valley. In the southernmost SETTING outcrops, up to 20 m red and green shale
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