Skeletonized Microfossils from the Lower–Middle Cambrian Transition of the Cantabrian Mountains, Northern Spain

Skeletonized microfossils from the Lower–Middle Cambrian transition of the Cantabrian Mountains, northern Spain SÉBASTIEN CLAUSEN and J. JAVIER ÁLVARO Clausen, S. and Álvaro, J.J. 2006. Skeletonized microfossils from the Lower–Middle Cambrian transition of the Cantabrian Mountains, northern Spain. Acta Palaeontologica Polonica 51 (2): 223–238. Two different assemblages of skeletonized microfossils are recorded in bioclastic shoals that cross the Lower–Middle Cambrian boundary in the Esla nappe, Cantabrian Mountains. The uppermost Lower Cambrian sedimentary rocks repre− sent a ramp with ooid−bioclastic shoals that allowed development of protected archaeocyathan−microbial reefs. The shoals yield abundant debris of tube−shelled microfossils, such as hyoliths and hyolithelminths (Torellella), and trilobites. The overlying erosive unconformity marks the disappearance of archaeocyaths and the Iberian Lower–Middle Cambrian boundary. A different assemblage occurs in the overlying glauconitic limestone associated with development of widespread low−relief bioclastic shoals. Their lowermost part is rich in hyoliths, hexactinellid, and heteractinid sponge spicules (Eiffelia), chancelloriid sclerites (at least six form species of Allonnia, Archiasterella, and Chancelloria), cambroclaves (Parazhijinites), probable eoconchariids (Cantabria labyrinthica gen. et sp. nov.), sclerites of uncertain af− finity (Holoplicatella margarita gen. et sp. nov.), echinoderm ossicles and trilobites. Although both bioclastic shoal com− plexes represent similar high−energy conditions, the unconformity at the Lower–Middle Cambrian boundary marks a drastic replacement of microfossil assemblages. This change may represent a real community replacement from hyolithelminth−phosphatic tubular shells to CES (chancelloriid−echinoderm−sponge) meadows. This replacement coin− cides with the immigration event based on trilobites previously reported across the boundary, although the partial infor− mation available from originally carbonate skeletons is also affected by taphonomic bias. Key words: Skeletonized microfossils, cambroclavids, chancelloriids, hyoliths, hyoliththelminths, sponges, benthic re− placement, Cambrian, Cantabrian Mountains, Spain. Sébastien Clausen [Sebastien.Clausen@univ−lille1.fr] Laboratoire de Paléontologie et Paléogéographie du Paléo− zoïque, UMR 8014 CNRS, Université des Sciences et Technologies de Lille, UFR Sciences de la Terre, 59655−Villeneuve d’Ascq, France; J. Javier Álvaro [Jose−Javier.Alvaro@univ−lille1.fr] Laboratoire de Paléontologie et Paléogéographie du Paléozoïque, UMR 8014 CNRS, Université des Sciences et Technologies de Lille, UFR Sciences de la Terre, 59655−Villeneuve d’Ascq, France; and Departamento Ciencias de la Tierra, Universidad de Zaragoza, 50009−Zaragoza, Spain. Introduction of their particular paleoecological and environmental condi− tions (Álvaro et al. 1999, 2003; Álvaro and Clausen in press). The Lower–Middle Cambrian transition, in its regional Medi− The Lower–Middle Cambrian boundary, defined in the terranean sense (see recent summary in Álvaro and Clausen Iberian Peninsula by the lowest appearance of the trilobite 2005), is a key interval for testing the relationship between Acadoparadoxides mureroensis (Sdzuy, 1958), is linked in the stratigraphic completeness of the fossil record and the the Iberian Chains (NE Spain) with the Valdemiedes event, a benthic community replacements recorded in different parts synchronous change in fauna related to a stepwise immigra− of the western Gondwana margin. This time span was rich in tion of relatively cosmopolitan invaders (Álvaro et al. 1999). paleogeographic and paleoecological events in West Gond− At present day, the stratigraphic successions in which this wana, such as the stepwise decrease in carbonate produc− event is recognized lie exclusively in the Iberian Chains. The tivity, the abrupt disappearance of their associated microbial occurrence of A. mureroensis in the Cantabrian Mountains and archaeocyathan−microbial reef ecosystems, the record of was reported by Álvaro et al. (1993) in the Esla nappe, a region an episodic tectonic instability correlatable across different recently analyzed from sedimentologic and sequence strati− platforms, and a drastic change to open−sea conditions that graphic viewpoints (Álvaro et al. 2000b). The importance of favored the appearance of a relatively cosmopolitan trilobite the Lower–Middle Cambrian transition in the Esla nappe is re− fauna (see recent summaries in Álvaro et al. 2000a, 2003; lated to two distinct paleogeographic patterns: the occurrence Álvaro and Vennin 2001). However, each platform recorded of archaeocyathan−microbial buildups (Debrenne and Zamar− different benthic community replacements as a consequence reño 1970; Perejón and Moreno−Eiris 2003) representing the Acta Palaeontol. Pol. 51 (2): 223–238, 2006 http://app.pan.pl/acta51/app51−223.pdf 224 ACTA PALAEONTOLOGICA POLONICA 51 (2), 2006 Fig. 1. A. Pre−Hercynian outcrops of the Iberian Peninsula (stippled) and setting of the Esla Nappe in the Cantabrian Zone (grey area). B. Geologic map of the Esla nappe and stratigraphic sections sampled in this work (Cr1–4 and V1; after Truyols et al. 1990; Aramburu et al. 1992; Álvaro et al. 2000b). youngest Lower Cambrian archaeocyathan zone of the Iberian this member is composed of grey limestone with abundant Peninsula (Moreno−Eiris et al. 1995), and the presence of a re− birdseye fabric and archaeocyathan−microbial reefs (Debren− gional unconformity within the Láncara Formation at the ne and Zamarreño 1970), and ooid−bioclastic limestone (Ál− Lower–Middle Cambrian boundary (Mohr 1969; Aramburu et varo et al. 2000b). The base of the Middle Cambrian is lo− al. 1992; Aramburu and García Ramos 1993). cated at the base of the upper member (with 1–40 m of global The present work establishes the taxonomic basis of the thickness), which consists of grey and pink, ooid−bioclastic skeletonized microfossils (“small shelly fossils”) for pur− limestone rich in glauconite (“Beleño facies”), overlain by an poses of paleoecological interpretation of the benthic com− alternation of reddish to purple, nodular limestone and shale munity replacements recorded across the Lower–Middle that is 0–30 m thick (“griotte” or “Barrios facies” Zamarreño Cambrian transition in the Esla nappe. As West Gondwanan 1972). The unconformity at the Lower–Middle Cambrian skeletonized microfossils are reasonably sparse, the descrip− boundary in the Esla nappe (Fig. 2), identified at the contact tion of these phosphatized microfossils will increase the between the two members, is associated with the abrupt dis− biogeographic discussions currently based on trilobites (Ál− appearance of reef mounds and their associated archaeo− varo et al. 2003). cyaths and the rapid immigration of diverse, lowermost Mid− dle Cambrian, relatively cosmopolitan trilobites. This hiatus Institutional abbreviation.—DGO, Faculty of Geology of is associated with a basinward facies shift capped by a major the Oviedo University, Oviedo, Spain. unconformity separating sharp changes in the type of carbo− nate deposition (Álvaro et al. 2000b). Neither C isotope ra− tios nor best preserved Sr isotope ratios show any major Geologic setting and stratigraphy change at the precise level of the boundary that might indi− cate a significant depositional hiatus (Shields et al. 2004). The fossils described herein occur in the Cantabrian Zone, After a period of erosion, sedimentation continued in a Mid− which is a major tectonostratigraphic unit of the Variscan dle Cambrian transgressive trend in which Lower Cambrian foreland fold belt located in the northwest of the Iberian Pen− sediments are found as reworked clasts in breccia layers. insula (Fig. 1). In this area, the Lower–Middle Cambrian transition lies in the Láncara Formation, which was subdi− vided by Zamarreño (1972) into two members. The lower Material and methods member (100–225 m thick) is composed of yellow dolo− stone, commonly rich in ooids, peloids and stromatolites. In The microfossils reported here were found up to 1 m below some areas (such as the Esla nappe), the uppermost part of and above the unconformity at the Iberian Lower–Middle CLAUSEN AND ÁLVARO—CAMBRIAN MICROFOSSILS FROM SPAIN 225 2m 1 0 Fig. 2. Stratigraphic units of the Lower–Middle Cambrian boundary interval in the Esla nappe, and setting of the fossils described in the text. Cambrian boundary. Samples were taken within the upper− Material.—Ten specimens preserved as secondarily phos− most Lower Cambrian ooid−bioclastic limestone, and from the phatized spicules in the Middle Cambrian “Beleño” facies, lowermost Middle Cambrian “Beleño” facies of the Láncara sections Cr2 and Cr4. Formation in the vicinity of Valdoré and Crémenes (V1 and Description.—The spicules have a regular six−rayed disk, Cr1–4 sections, respectively; Fig. 1). Thirty kilograms of lime− 310–330 µm in diameter, with rays diverging at ca. 60°; the stone were etched with 10% acetic acid. Chemical composi− rays are slender and can be of unequal length (80–200 µm); tion of extracted microfossils was determined using SEM− they usually intersect at a slight angle to the plane of the cen− EDS analyser. Although some of the extracted microfossils tral disk so that both concave and convex sides are distin− were originally phosphatic in composition, the phosphate is guishable; lack of central transverse ray. commonly the result of diagenetic replacement of the whole skeletons, superficial impregnation, or steinkerns

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