J.C. Gutiérrez-Marco, I. Rábano and D. García-Bellido (eds.), of the World. Cuadernos del Museo Geominero, 14. Instituto Geológico y Minero de España, Madrid. ISBN 978-84-7840-857-3 © Instituto Geológico y Minero de España 2011

BIOSTRATIGRAPHY OF THE MIDDLE ORDOVICIAN FROM CENTRAL SPAIN

J. Reyes-Abril1, J.C. Gutiérrez-Marco2 and E. Villas3

1 Escuela de Ingeniería Geológica, Grupo TERRA, Universidad de Los Andes, Núcleo Pedro Rincón Gutiérrez s/n, 5101 Mérida, Venezuela. [email protected] 2 Instituto de Geociencias (CSIC-UCM), Facultad CC. Geológicas, José Antonio Nováis 2, 28040 Madrid, Spain. [email protected] 3 Departamento de Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Pedro Cerbuna s/n, 50009 Zaragoza, Spain. [email protected]

Keywords: Biostratigraphy, Rhynchonelliformean brachiopods, Central Iberian Zone, Darriwilian.

INTRODUCTION

A recent taxonomic study on the rhynchonelliformean brachiopods from the Darriwilian dark shales of the Central-Iberian Zone, Central Spain (Reyes-Abril, 2009; Reyes-Abril et al., 2010) has considerably increased the number of orthides and strophomenides known across the whole Iberian Peninsula. The studied brachiopods were collected in 58 localities of 6 provinces from the regions of Castilla-La Mancha, Andalucía and Extremadura. Middle Ordovician brachiopods previously known in the area were mainly derived from upper Darriwilian rocks, while most of the new studied brachiopods are from middle Darriwilian rocks. A total of 21 genera are now known from the whole Darriwilian strata of Iberia including the strophomenide genera Aegiromena and Dactylogonia, as well as 19 orthide genera, including Almadenorthis, Gutiorthis, Orthambonites, Paralenorthis, Sivorthis, Apollonorthis, Atlantida, Brandysia, Eodalmanella, Howellites, Cacemia, Heterorthina, Tissintia, Mcewanella, Crozonorthis, Nocturnellia and Lipanorthis, plus two new genera of the families Cremnorthidae and Harknessellidae. Considering the regional biostratigraphical and biochonological context, in which the Ibero-Armorican biozones can be easily correlated with biozones based on other fossil groups, we have followed the Mediterranean stratigraphic scale, originally proposed in Bohemia (Havlícˇek and Marek, 1973) and updated and completed in Iberia (Gutiérrez-Marco et al., 1995, 2002). Thus we are referring the stratigraphic range of the studied brachiopods to the Mediterranean regional stages Oretanian and Dobrotivian, approximately correlatable to the middle and upper stage slices (Dw2-3) of the global Darriwilian stage. Exception made with the uppermost Dobrotivian, which extends into the Upper Ordovician, correlating with the lowermost Sandbian (Gutiérrez-Marco et al., 2008; Bergström et al., 2009). Brachiopods were first used in the late nineteenth century to subdivide the Ordovician succession in Iberia (Delgado, 1897). Since then, different brachiopod biozones have been proposed for the Middle Ordovician rocks of Spain (Born, 1918; García Alcalde and Arbizu,1982; Villas, 1985; Young, 1985; Gutiérrez-Marco et al., 1984, 2002). Recent improvements in the detailed stratigraphical knowledge of

463 J. Reyes-Abril, J.C. Gutiérrez-Marco and E. Villas many successions in the Central Iberian Zone, and on the taxonomy of important brachiopod species, have allowed the redefinition and updating of previous brachiopod biozones, which are examined in the present note.

Figure 1. Stratigraphic correlation of the proposed Middle Ordovician brachiopod biozones, with the vertical range of known brachiopod species from Central Spain.

STUDIED BIOZONES

The stratigraphic range of the 27 brachiopod species identified in a wide area of the southern Central Iberian Zone (Reyes-Abril, 2009; Reyes-Abril et al., 2010) has allowed to review the biostratigraphical units defined upon brachiopods by previous authors. Our contribution recognizes, from base to top, the following four biozones: 1, Orthambonites–Sivorthis noctilio Partial Range Biozone (lower Oretanian; base of the Darriwilian 2); 2, Cacemia ribeiroi Taxon Range Biozone (upper Oretanian; upper Darriwilian 2); 3, Heterorthina morgatensis Taxon Range Biozone (uppermost Oretanian to lower Dobrotivian; lower Darriwilian 3), and 4, Heterorthina kerfornei–Aegiromena mariana Partial Range Biozone (lower to lower

464 BIOSTRATIGRAPHY OF THE MIDDLE ORDOVICIAN BRACHIOPODS FROM CENTRAL SPAIN upper Dobrotivian; uppermost Darriwilian 3). Although a pre-Oretanian species, Nocturnellia praedux (Havlícˇek in Arbin et al., 1978), occurs below the base of the lowest of the four biozones considered, its extreme rarity and its scattered occurrence in the lower Oretanian precludes its consideration as a biostratigraphic marker. The regional brachiopod biozonation can be applied to the Portuguese part of the Central Iberian Zone, but not to areas in northern Spain, where the lower and upper Dobrotivian assemblages include some taxa not recovered from Central Iberia. Some common brachiopods in the Middle Ordovician of the studied area, including the index-species of the proposed biozones, are illustrated in Plate 1.

Orthambonites–Sivorthis noctilio Biozone

This is a Partial Range Biozone, with the base defined by the lowest record of the genus Orthambonites on a wide area of the Central Iberian Zone, mostly coinciding in vertical range with the genus Sivorthis, which appears slightly higher in the succession. The top of the biozone is marked by the disappearance of Sivorthis noctilio (Pl. 1: G), the species most abundant, ubiquitous and easily recognizable in the whole Ibero-Armorican area. Among the associated species exclusive to the biozone are Sivorthis calatravaensis, Paralenorthis estenaensis, Paralenorthis lolae, Gutiorthis incurvata, a new genus and species of the family Cremnorthidae, as well as a new species of Dactylogonia. Paralenorthis alata, a species known from the British Arenigian (Bates, 1969), presents here its youngest occurrence. Punctual records are also known of Nocturnellia praedux, Eodalmanella sp. and Lipanorthis sp. The occurrence of Almadenorthis auriculata, restricted originally to its type locality, is now found below the base of the biozone, although it could be extended up through it in future studies. The upper boundary of the biozone coincides with a barren interzone for rhynchonelliformean brachiopods, which precedes the first record of the nominal form of the overlying biozone. The Orthambonites–S. noctilio Biozone spreads through the southern part of the Central Iberian Zone (lower beds of the Navas de Estena Shales and the Navatrasierra Shales; lower and middle beds of the Río Shales), as well as through the middle of the Valongo and Moncorvo formations in Portugal (Sá, 2005). In the Armorican Massif (France) the biozone can also be recognized in the lower part of the Traveusot Formation in the synclines south of Rennes (Tromelin and Lebesconte, 1876; Pillet et al., 1990), where S. noctilio an the same new genus and species of the family Cremnorthidae are also represented. The biozone proposed herein substitutes in range and meaning the old “Monorthis” noctilio or “Orthis” noctilio Biozone, considered by Gutiérrez-Marco et al. (1984, 2002) and San José et al. (1992), after the taxonomic revision of the nominal species and the redefinition of its base. This way, the first record of such regionally characteristic genera as Orthambonites, Paralenorthis or Gutiorthis, are included in the Biozone. Considering the brachiopods represented within the association, the biozone is also equivalent to the upper part of the “Didymograptus Shales” and to the “Orthis noctilio Shales” of Delgado (1908). The characteristic brachiopods of the “Orthis calligramma Zone” (Born, 1918) and the “Hesperorthis Biozone” (García Alcalde and Arbizu,1982) occur also in the revised biozone. The occurrence of some index fossils of the biozone such as “Orthis” noctilio and “Orthis” miniensis in the Upper Ordovician of Sardinia (Meneghini, 1857; Vinassa de Regny, 1927; Leone, 1998) is unlikely and both were probably confused with younger forms. In this sense, and according to Havlícˇek et al. (1987), the Sardinian specimens identified as O. noctilio, O. noctilio novata, and even as O. calligramma by Vinassa de Regny (1927), really belong to Nicolella actoniae, a characteristic species for the late Katian (Upper Ordovician) of the Mediterranean Region.

465 J. Reyes-Abril, J.C. Gutiérrez-Marco and E. Villas

From a chonostratigraphic point of view, the dating of the Orthambonites–S. noctilio Biozone as lower Oretanian, made by Gutiérrez-Marco et al. (1984, 2002), is corroborated with the record of graptolites from the Didymograptus artus Biozone throughout the whole range of the unit. The lower Oretanian is correlated to the first half of the Darriwilian 2 of the Global Scale.

Cacemia ribeiroi Biozone

This biozone coincides with the vertical range of Cacemia ribeiroi, besides which, only a new species of Heterorthina occurs in its uppermost beds. The unit is widely spread through the whole Ibero-Armorican area, where the record of the graptolite Didymograptus murchisoni, which pre- and postdates this biozone, indicates an late Oretanian age (latest Darriwilian 2 substage). Cacemia ribeiroi is a frequent and easily identifiable species, because its auriculated outline and relatively fine costellation (Pl. 1: M). Several authors have pointed out its stratigraphic value and used it to propose the “Orthis Ribeiroi Shales” in Bussaco (Delgado 1897, 1908), the “Orthis ribeiroi Zone” (Born, 1918) and the “Cacemia Biozone” (García-Alcalde and Arbizu, 1982) in Almadén, and the “Cacemia ribeiroi Biozone” in the southern Central Iberian Zone (Gutiérrez-Marco et al., 1994, 2002). However, mixed collections have been detected for the establishment of some of these units in the works of Delgado(1908) and Born (1918), with specimens coming from younger beds but also, occasionally, from lower strata, such as when C. ribeiroi was confused with other auriculated brachiopods from the lower Oretanian (Sivorthis, Paralenorthis). This is also the case of “Orthis vespertilio” Sow., a species repeatedly confused with C. ribeiroi in the Central Iberian Zone, following an erroneus identification and illustration by Verneuil and Barrande (1855) and Mallada (1875), in which part of the citations correspond with lower Oretanian localities. The C. ribeiroi Biozone has been recognized in the Navas de Estena and Navatrasierra formations and in the upper part of the Río shales of the studied area. It is also represented in the Luarca Shales of the West Asturian-Leonese Zone (Gutiérrez-Marco et al., 1999). Outside Spain, the nominal species has been described from the upper part of the Brejo Fundeiro Formation (Cacemes Group) of Bussaco (Portugal), as well as in the lower part of the Postolonnec Formation of the Armorican Massif, France (Mélou, 1976). It has been also identifed, with doubts, in the Algerian Sahara (Mélou et al., 1999). Above the C. ribeiroi Biozone, an upper Oretanian graptolitic interval was recorded, yielding scarce trilobites and molluscs of broad vertical range. Nevertheless, in a Montes de Toledo section, these beds yielded a new species of the plectorthid Atlantida, a genus so far restricted to the upper Darriwilian of Morocco (Havlícˇek, 1971) and, in consequence, of little biostratigraphic value.

Plate 1. Some Middle Ordovician brachiopods from the Central Iberian Zone, including the index species of the proposed brachiopod biozones. Scale bars 5 mm, except where otherwise indicated. A, Aegiromena mariana Drot, 1969, ventral internal mould, MGM- 6477-O, scale bar 2 mm, Calzada de Calatrava. B, Dactylogonia asturica (Villas, 1989), dorsal internal mould, MGM-6453-O, La Alameda. C, Gutiorthis incurvata Reyes-Abril and Villas (in Reyes-Abril et al., 2010), ventral internal mould, MGM-5968-O, Navas de Estena. D-E, Orthambonites sp., ventral internal mould, MGM-5999-O, Navas de Estena (D) and latex cast of dorsal exterior, MGM- 6007-O, Ventas con Peña Aguilera (E). F, Paralenorthis estenaensis Reyes-Abril and Villas (in Reyes-Abril et al., 2010), ventral internal mould, MGM-6072-O, Navas de Estena. G, Sivorthis noctilio (Sharpe, 1849), ventral internal mould, MGM-6249-O, Ventas con Peña Aguilera. H, Heterorthina morgatensis Mélou, 1975, ventral internal mould, MGM-6792-O, Retuerta del Bullaque. I, L, Heterorthina kerfornei Mélou, 1975, dorsal internal mould, MGM-6758-O, Calzada de Calatrava (I) and ventral internal mould, MGM-6729-O, Calzada de Calatrava (L). J, Apollonorthis bussacensis (Sharpe in Ribeiro et al., 1853), ventral internal mould, MGM-6509-O, Calzada de Calatrava. K, Nocturnellia praedux (Havlícˇek in Arbin et al., 1978), internal moulds of two ventral valves, (left) MGM-6937-O, (right) MGM-6938-O, scale bar 2mm, Solana del Pino. M, Cacemia ribeiroi (Sharpe in Ribeiro et al., 1853), ventral internal mould, MGM-6335-O, Helechosa de Los Montes. N, Crozonorthis musculosa Mélou, 1976, ventral internal mould, MGM-6988-O, Calzada de Calatrava.

466 BIOSTRATIGRAPHY OF THE MIDDLE ORDOVICIAN BRACHIOPODS FROM CENTRAL SPAIN

467 J. Reyes-Abril, J.C. Gutiérrez-Marco and E. Villas

Heterorthina morgatensis Biozone

It is defined by the vertical range of its nominal species, and thus coincides with the H. morgatensis Biozone proposed by Villas (1985) in the Iberian Chains (NE Spain) and by Young (1985) in the Serra do Bussaco (Portugal). Besides H. morgatensis, also Crozonorthis musculosa, Aegiromena mariana and an undetermined dalmanellidine occur in the upper beds of the biozone. In its lower beds a new species of Eodalmanella has been recorded (Reyes-Abril, 2009). The H. morgatensis Biozone is widely recognized throughout the Ibero-Armorican region, having been identified in the middle part of the Navas the Estena and Navatrasierra Shales, as well as in the El Caño Alternation of the Central Iberian Zone. Within the Navatrasierra Shales, common records of H. morgatensis lie below and above the laterally discontinuous Los Rasos Sandstones Member, being sometimes recorded in coquinoid beds within these sandy tempestites, or in its Eastern Sierra Morena equivalent El Caño Alternation (Mélou, 1975). In the Portuguese extension of the Central Iberian Zone, the H. morgatensis Biozone can be identified in Penha Garcia and in the Serra do Bussaco, at the top of Brejo Fundeiro Formation and in the base of the Fonte da Horta Formation (Henry et al., 1976; Young, 1985). It has also been recognized at the Iberian Chains (Villas, 1985) and the Cantabrian Zone of the Iberian Massif (Gutiérrez-Marco et al., 1996, 1999; Gutiérrez-Marco and Bernárdez, 2003). In the Armorican Massif, H. morgatensis is typically recorded in the Postolonnec Formation, within the shales that overlie the Kerarvail Sandstones, a local equivalent to the Los Rasos and Monte da Sombadeira formations (Mélou, 1975; Henry et al., 1976), as well as in the sandstone beds at the base of the Mont de Besneville Formation, in Normandy. The record of the species by Mélou (1975) in another Armorican locality, Andouillé-La Touche, at the top of the Andouillé Formation, is very unlikely. Those beds correspond to the trilobite biozone of Placoparia borni (Subzone of Marrolithus bureaui), which has been dated with graptolites and chitinozoans as lowermost Sandbian (Upper Ordovician). The form identified there by Mélou (1975) may coincide with another species from La Touche, determined as Heterorthina sp. by Young (1985: Pl. 21, figs. 8-15), but that could belong to a different genera like those occurring on a similar stratigraphic position at the top of the Postolonnec Formation (Botquelen and Mélou, 2007). Chronostratigraphycally, the base of the H. morgatensis Biozone can be referred to the Upper Oretanian based on the occurrence of graptolites from the Didymograptus murchisoni Biozone at the Cantabrian Zone and in a few Central Iberian localities (Gutiérrez-Marco et al., 1994, 1999, 2002). Nevertheless, most of its development corresponds to the lower Dobrotivian, and the biozone can be correlated, in a broad sense, with the lower half of the Darriwilian 3 substage.

Heterorthina kerfornei–Aegiromena mariana Biozone

This Partial Range Biozone is characterized by the almost total concurrent range of Heterorthina kerfornei and Aegiromena mariana. The base of the biozone coincides with the acme of Aegiromena mariana, which frequently crowds many bedding planes. In Central Iberia, the lower part of the biozone can be correlated with the Morgatia hupei trilobite Subzone, where the brachiopod assemblage reaches its highest diversity. Besides the two nominal species, Howellites hammanni and Harknesellidae gen. et sp. nov. occur frequently, in addition to the local acme of Crozonorthis musculosa and the earliest record of Apollonorthis bussacensis. In higher beds of the unit, yet coinciding with the Placoparia (Coplacoparia) borni trilobite biozone, the brachiopod association is dominated by H. kerfornei and A. mariana, with the appreciable decrease of C. musculosa (restricted to a few horizons), and the sporadic record of

468 BIOSTRATIGRAPHY OF THE MIDDLE ORDOVICIAN BRACHIOPODS FROM CENTRAL SPAIN

Dactylogonia asturica or Apollonorthis bussacensis. The disappearance of Heterorthina kerfornei, slightly higher than that of A. mariana, marks the biozone top, which coincides with the lower boundary of the Lagenochitina ponceti chitinozoan biozone, but not with the disappearance of the trilobite Neseuretus tristani. The re-definition of the H. kerfornei–A. mariana Biozone (name adapted from Gutiérrez-Marco et al., 2002) is intended to solve the identification problems of other Dobrotivian brachiopod biozones overlying the H. morgatensis Biozone. Different solutions have been given by other authors to the difficulty of evaluating the biostratigraphic meaning of local occurrences of very conspicuous species such as C. musculosa or A. bussacensis. This is especially complicated in the absence of an effective biostratigraphic control provided by graptolites, chitinozoans or trilobites. The H. kerfornei–A. mariana Biozone, such as it is considered in this study, is a stratigraphical equivalent of the A. mariana and E. musculosa biozones of García-Alcalde and Arbizu (1982), and also equates the combination of the A. mariana–E. musculosa Concurrent Range Biozone and the H. kerfornei Taxon Range Biozone defined by Villas (1985). It is also coincident with the H. kerfornei Partial Range Biozone plus the ranges of those brachiopods occurring in the Placoparia (Coplacoparia) borni Taxon Range Biozone, as considered by Young (1985). The unit also includes the H. kerfornei Biozone of Gutiérrez-Marco et al. (1984, 1995) and San José et al. (1992). In the study area, the H. kerfornei–A. mariana Biozone is represented in the upper part of the Navas de Estena Shales and in the Navatrasierra Shales; it also typically occurs in the Guindo Shales and in the lower part of the Botella Quartzite of the southernmost part of the Spanish Central Iberian Zone. In the Portuguese part, the biozone extends through most of the Fonte da Horta Formation, the Cabril Formation and probably, also, through part of the Carregueira Formation at Bussaco, Penha Garcia and Dornes (Henry et al., 1976; Young, 1985). The base of the unit could be represented in the upper third of the Valongo Formation of the Valongo-Arouca region (Couto et al., 1997), as well as in the Moncorvo Formation of Trás- os Montes (Sá, 2005), although detailed taxonomic studies are needed to verify it. Different occurrences of the biozone at the Iberian Peninsula are also known from the Iberian Chains (Villas, 1985) and the Cantabrian Zone (Truyols et al., 1996; Gutiérrez-Marco et al., 1999). Outside the Iberian Peninsula, the nominal brachiopods are recorded in different localities from the Andouillé, Postolonnec and Travesout formations of the Armorican Massif (Mélou, 1973, 1975), with the exception of A. mariana in the Postolonnec Formation. Occasionally, the biozone can be characterized in North Africa, according to the study by Mélou et al. (1999) who record the co-occurrence of the two index species in Algeria, in beds yielding also A. bussacensis and Tenuiseptorthis niliensis. Depending on the region considered within the studied area, the upper part of the H. kerfornei–A. mariana Biozone can be incomplete, since the pelitic beds characterizing it change laterally in the upper Dobrotivian to alternations of sandstones and shales, culminating in massive quartzites (Retuerta Sandstones, La Cierva or Botella Quartzites). In these coarse-grained units, corresponding to shallow and turbulent environments, the records of the nominal species are interrupted. The only brachiopods occurring there are forms restricted to sandy facies, particularly Tissintia immatura, a species geographically widespread that also occurs in older units (Los Rasos Sandstones). The reference by Mélou (1975) to the occurrence of H. morgatensis in the Botella Quartzite at Sierra Morena, could actually correspond also to Tissintia. T. immatura is also known from the Dobrotivian sandstones of the Sierra de San Pedro (Elice Formation). However, Tissintia is not the only brachiopod occurring in the sandy facies from the Iberian Dobrotivian, since Tafilaltia has also been cited from the top of the Retuerta Sandstones at the Toledo Mountains (Montero, 1989) and with certainty it is yielded by correlative sandstones at the Hesperian Chains of the Iberian Cordillera. This is why San José et al. (1992) and Gutiérrez-Marco et al. (2002) came

469 J. Reyes-Abril, J.C. Gutiérrez-Marco and E. Villas to propose the erection of a Tafilaltia Sub-biozone within the uppermost Dobrotivian sandstones. The base of the H. kerfornei–A. mariana biozone is placed within the Gondwanan range of the Hustedograptus teretiusculus graptolite biozone, and its top lies within the Oepikograptus bekkeri Biozone, paralleled by the Nemagraptus gracilis Biochronozone. Therefore, its total range extends from the high- lower Dobrotivian to the terminal-upper Dobrotivian, correlatable in global terms to the upper half of the Darriwilian 3 and the basal beds of the Sandbian 1, bridging the boundary between the Middle and the Upper Ordovician series. However, most of the Central Iberian fossiliferous localities included in this biozone are from the uppermost Darriwilian, and have not been detected yet within the sandy and quartzitic facies that locally characterize the lowermost Sandbian.

Acknowledgements

This is a contribution to the project CGL2009-09583 of the Spanish Ministry of Science and Innovation. Diego García-Bellido (CSIC, Madrid) is thanked for revising the English version of this paper.

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