LATE MOSCOVIAN TO EARLY KASIMOVIAN FUSULINES FROM THE ÁNDARA MASSIF, PICOS DE EUROPA (PENNSYLVANIAN, CANTABRIAN ZONE, NORTHERN SPAIN)

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1 Journal of Foraminiferal Research, v. 45, no. 3, p , July 2015 LATE MOSCOVIAN TO EARLY KASIMOVIAN FUSULINES FROM THE ÁNDARA MASSIF, PICOS DE EUROPA (PENNSYLVANIAN, CANTABRIAN ZONE, NORTHERN SPAIN) ELISA VILLA *,OSCAR MERINO-TOMÉ AND JUAN R. BAHAMONDE Departamento de Geología, Universidad de Oviedo, Oviedo, Spain ABSTRACT Extensive outcrops of fusuline-rich limestone strata of Pennsylvanian age occur in the Eastern or Ándara Massif of the Picos de Europa mountains in Northern Spain. The stratigraphic section is composed of the upper part of the Picos de Europa Formation, a carbonate succession deposited on a broad platform top in a distal foreland-basin margin setting, and the unconformable overlying Las Llacerias Formation, which represents carbonate-ramp strata deposited during the deformation of the underlying Picos de Europa platform. The tectonic setting allowed the accumulation of a very thick stratigraphic succession during the late Pennsylvanian, when the basin was located at paleoequatorial latitudes in the gulf of the western Palaeo-Tethys Ocean, adjacent to the eastern coast of Pangea. An in-depth stratigraphic and micropaleontological study involving six sections and several isolated localities revealed an interval spanning the uppermost Moscovian to the lowermiddle Kasimovian. This study has allowed us to establish a composite section, more than 400 m thick, recording the uppermost part of the Fusulinella Zone (F. alvaradoi subzone), the Protriticites Zone and the Montiparus Zone. Forty-one fusuline taxa are described, including four new species, Protriticites grajalensis, P. quasiovoidalis, P. benshae, and Pseudotriticites cantabricus. The composition of the fusuline assemblages differs from those of the Carboniferous in key areas of Eastern Europe, and reveals their affinity to Central Asian assemblages as part of a separate biogeographic province. In spite of these paleobiogeographic differences, the observations on the fusuline evolutionary lineages, as well as the presence of a few common forms, made it possible to establish an approximate correlation between the Ándara and the Eastern European successions. We also report a previously unknown thrust fault, refine the stratigraphy and age of the successions outcropping in the Ándara Massif, and determine the diachronism of the base of the unconformable Las Llacerias Formation. INTRODUCTION During the early 1990s, the IUGS Subcommission on Carboniferous Stratigraphy (SCCS) initiated a program devoted to the characterization of the Pennsylvanian stage boundaries. One of these levels, the Moscovian/Kasimovian boundary, lies within the stratigraphic interval examined in this paper. Although considerable progress has been made since then, neither a boundary marker nor a GSSP (Global Boundary Stratigraphic Section and Point) has yet been * Correspondence author. evilla@geol.uniovi.es proposed. The main difficulties in defining biostratigraphic markers for the boundary derive from the relatively strong provincialism observed in late Pennsylvanian biota. In the paleocircumequatorial areas, the Pennsylvanian fossil groups with the most potential for global correlation are conodonts and fusulines. A conodont, Idiognatohodus simulator (Ellison), has already been selected to mark the lower Gzhelian boundary (Heckel et al., 2008; Villa et al., 2009a). Additionally, I. saggitalis Kozitskaya in Kozitskaya et al., 1978 and I. turbatus Rosscoe & Barrick, 2009, are presently being discussed as possible markers for the lower Kasimovian boundary (Ueno and Task Group, 2014). If accepted, the latter would be placed at a level (close to the base of the Khamovnikian) younger than the present lower Kasimovian boundary of the Russian stratigraphy, which lies at the base of the Krevyakinian. A fusuline event (first appearance of the genus Protriticites) has also been proposed to serve as a marker for this boundary in its classical position (Davydov, 2007). In any case, whichever marker is eventually selected, a thorough knowledge of the fusulines around the proposed levels will be essential to further incorporate the marine Pennsylvanian successions from around the world into the biostratigraphic scheme emerging from these studies. The importance of fusulines for correlation derives not only from the relative abundance of these foraminifera in shallow carbonate platform strata, and the easy application to biostratigraphy of their evolutionary trends, but also from the huge amount of data compiled after more than a century of fusuline studies in relevant Carboniferous strata. Therefore, fusuline-bearing successions exhibiting a comparatively complete stratigraphic record may provide data of prime importance in this aim. The Picos de Europa region, situated in the eastern part of the Cantabrian Zone (Lotze, 1945) (Fig. 1), contains one of the most complete records of Carboniferous marine carbonates in the world. These sediments were deposited on the east coast of Pangea, within the western part of a gulf formed at paleoequatorial latitude by the Paleo-Tethys Ocean. A fossiliferous succession is exposed, recording from the very base of the Carboniferous to the Gzhelian stage. It consists of: a) Bashkirian to early Kasimovian carbonate sediments (up to 1500 m thick) deposited in the distal margin of a foreland basin and interpreted as a large, high-rising, microbial-dominated platform (Bahamonde et al., 2007); and b) a set of unconformable stratigraphic units, late Myachkovian to Gzhelian in age, that accumulated synchronously with the end of the Variscan Orogeny (Merino-Tomé et al., 2006, 2009a, b). This paper focuses on the upper Moscovian and lower to middle Kasimovian strata exposed in the Ándara Massif of the Picos de Europa. Six different sections have been 264 Journal of Foraminiferal Research fora d 14/7/15 13:19:23 264

2 FUSULINES FROM THE PICOS DE EUROPA 265 FIGURE 1. Sketch of the Cantabrian Zone showing the location of the Ándara Massif and the Las Llacerias section in the Picos de Europa mountains. studied and integrated into a composite section nearly 400 m thick that contains a diverse marine biota. Fusulines are the most conspicuous and well-represented fossils, but conodonts, corals, sponges, brachiopods, and algae are also present. Co-occurrence of fusulines and conodonts (Sanz- González in Villa et al., 2009b) makes this succession particularly interesting for the SCCS program devoted to global correlation. STRATIGRAPHY OF THE PENNSYLVANIAN CARBONATE SUCCESSION IN THE PICOS DE EUROPA REGION Lying conformably over the dark lime mudstones of the Barcaliente Formation (Serpukhovian), the microbialdominated carbonate platform successions exposed in the Picos de Europa region are represented by the Valdeteja and the Picos de Europa formations, the former essentially Bashkirian in age and the latter ranging from early Moscovian to earliest Kasimovian (Fig. 2). Seismic-scale exposures of the western carbonate platform margin show three different stratal domains: basin/toe-of-slope, slope and platform-top, and the clear angular-bedding relationship between them (Bahamonde et al., 2000, 2007). Fusulines are particularly abundant in the platform-top strata, which show very high sedimentation rates during late Myachkovian-early Krevyakinian times, recording the increase in accommodation space. This younger part (lower Protriticites Zone, Fig. 2) of the carbonate platform succession is present in the central and southern areas of the Picos de Europa region and shows stratigraphic features such as red, nodular-carbonate beds, well developed lateritic crusts and an increase in biotic diversity. Unconformable synorogenic carbonates of latest Myachkovian to Ghzelian age can be found overlying the Barcaliente, Valdeteja or Picos de Europa formations at different localities in the Picos de Europa region (Merino- Tomé et al., 2006, 2009a, b). In the central and southern sectors, synorogenic late Krevyakinian-Khamovnikian carbonate successions are assigned to the Las Llacerias Formation, which was described by Martínez-García & Villa (1999) in the Las Llacerias section. This lithostratigraphic unit consists of a dark-coloured, marly calcareous succession, in some localities reaching up to 190 m in thickness (Merino-Tomé et al., 2009a). In contrast with the Picos de Europa Formation, the Las Llacerias Formation shows significant changes in textures, microfacies and facies distribution, interpreted as being deposited in ramp-like carbonate systems. These carbonate ramps nucleated above elongated tectonic uplifts approaching sea-level that developed during the tectonic thrusting of previous carbonate successions (Merino-Tomé et al., 2009a, b). THE UPPER MOSCOVIAN-KASIMOVIAN CARBONATE SUCCESSION IN THE ÁNDARA MASSIF The part of the Carboniferous succession studied in this paper outcrops in the Ándara Massif along two stacked thrust units separated by a thrust fault (Fig. 3). The succession is composed of platform-top carbonates of the Picos de Europa Formation and the overlying Las Llacerias Formation, which contains numerous minor unconformities. In the lower thrust unit, the thrust fault defines a ramp surface cutting, from north to south, strata of the Las Llacerias Formation at progressively higher stratigraphic levels. In the upper thrust unit, the detachment surface almost follows the same stratigraphic interval within the platform-top strata of the Picos de Europa Formation (Fig. 3). The complete stratigraphy of the carbonate succession cannot be reconstructed on the basis of a single stratigraphic section because the angular unconformity at the base of the Las Llacerias Formation truncates the top of the Picos de Europa Formation at different stratigraphic levels. In addition, several younger unconformities, developed during the deposition of the Las Llacerias Formation, caused the erosion of older strata of this stratigraphic unit at some localitites. Six stratigraphic sections were logged (Fig. 4) and an intensive program of fusuline sampling was undertaken, including several additional localities scattered throughout the study area (Fig. 3). This study has it made possible: a) to reconstruct the detailed stratigraphy of the platform-top strata of the Picos de Europa and Las Llacerias formations, b) to document the fusuline biostratigraphy, c) to determine the extent of the gap involved in the unconformity separating the stratigraphic units (Fig. 5), and d) to ascertain the age of the base of the Las Llacerias Formation at different localities. Journal of Foraminiferal Research fora d 14/7/15 13:19:24 265

3 266 VILLA AND OTHERS FIGURE 2. Stratigraphy of the Pennsylvanian carbonate successions of the Picos de Europa area indicating the major stratal domains of the microbial carbonate platform of the Valdeteja and Picos de Europa formations. Main lithofacies associations and the large-scale platform geometries are indicated (modified from Bahamonde et al., 2007). Vertical bar: stratigraphic succession studied in this paper. Picos de Europa Formation The uppermost 300 m of the Picos de Europa Formation are exposed in the Mazarrasa (MZ), Morra de Lechugales peak (ML), Castillo del Grajal (CG) and Silla de Caballo Bajero (SCB) sections (Figs. 3 4). These strata consist of a cyclic succession of microbial/phylloid algal boundstones, reddish pseudonodular mud- to wackestones, and thinner packages of grain-supported shallow-water deposits capped by subaerial exposure surfaces. Remarkable red-matrix karstic breccias are associated with the most prominent subaerial-exposure surfaces. According to its fusulinoidean content, the part of the Picos de Europa Formation exposed at the Mazarrasa and Morra de Lechugales sections belongs to the Fusulinella alvaradoi subzone (uppermost part of the Fusulinella Zone) and is late Myachkovian in age. In the Castillo del Grajal section, the Picos de Europa Formation is characterized by the occurrence of primitive forms of Protriticites near the base of the section, and by more evolved species of this genus in younger beds. These younger beds are also recognized in the Silla de Caballo Bajero section. In spite of the great thickness of the Picos de Europa Formation, its top does not reach the top of the Protriticites Zone, which extends into the overlying Las Llacerias Formation (Figs. 4 6). Las Llacerias Formation This lithostratigraphic unit comprises at least three unconformity-bound sequences with a discontinuous spatial extention (Merino-Tomé et al., 2009a; Fig. 4). Sequences 1 and 2 belong in this sector to the upper part of the Protriticites Zone (late Krevyakinian-earliest Khamovnikian in age) and occur only in the lower thrust nappe. Both sequences are well exposed in the Picos del Jierro section. Sequence 1 reaches nearly 90 m in thickness and its base is a remarkable unconformity surface with karst features showing discontinuous calcareous breccia lenses. This surface is overlain by a 13 m-thick stratal package with remarkable lateral continuity, which can be traced from the Picos del Jierro section along the foothill of the Grajales Ridge (locality AND-8) up to the top of the Mazarrasa section (Figs. 3 4). It consists of dark, nodular, and very fossiliferous wacke- to packstones, oncoidal float- to rudstones with intercalated chaetetid- and syringoporidrich boundstones, and phylloid and Anthracoporella bafflestones. Overlying this basal interval, dark and massive bioclastic wackestones form three decametre-thick massive packages separated by well-bedded, metre-thick stratal packages of fossil-rich dark marls and black nodular wackestones. Fusulines from the youngest strata of Journal of Foraminiferal Research fora d 14/7/15 13:19:30 266

4 FUSULINES FROM THE PICOS DE EUROPA 267 FIGURE 3. A, 3D reconstruction showing the trace of the two structural units sampled. B, Geological map of the study area. CG: Castillo del Grajal section, MZ: Mazarrasa section, PJ: Picos del Jierro section, SCB: Silla de Caballo Bajero section, SCC: Silla de Caballo Cimero section, ML: Morra de Lechugales section, AND: localities along the Grajales Ridge. Journal of Foraminiferal Research fora d 14/7/15 13:19:45 267

5 268 VILLA AND OTHERS FIGURE 4. Correlation, stratigraphy and location of samples in the six studied sections. Sequence 1 in the Picos del Jierro section have already been studied (Maas, 1974; Maas & van Ginkel, 1986; van Ginkel & Villa, 1991; Martínez-Chacón et al., 2010), these beds being referred to as the Tabla de Lechugales mountain peak (van Ginkel & Villa, 1991) and to Morra de Lechugales section (Martínez-Chacón et al., 2010). Sequence 2 is only 25 m thick and its top is truncated by a thrust fault. The lower part of the sequence (approximately 8 m thick) consists of graded decimetre-to-centimetre thick tabular beds of detrital, intraclastic and skeletal rudstones, and grain- to packstones alternating with dark marls. The upper part (17 m thick) is composed of darkgray nodular mud- to wackestones, algal bafflestones and massive microbial boundstone. Strata of the Las Llacerias Formation yielding Montiparus species, which indicate a late Khamovnikian age, have been grouped into an unconformity-bound stratigraphic unit named Sequence 3. They overlie the Picos de Europa Formation in the Castillo del Grajal and Silla de Caballo Cimero sections and along the Grajales Ridge (Fig. 3), a mountain crest that links the peaks of La Rasa and Grajal de Arriba. The base of these younger strata of the Las Llacerias Formation consists of an irregular and very penetrative erosional surface that is overlain by calcareous breccia and calcareous conglomerates with common quartzite pebbles, followed by dark gray, marly siltstones and shales with plant fragments. These deposits are, in turn, overlain by dark-nodular, fossiliferous wackestones and pack- to grainstones with intercalated phylloid and dasycladacean algal bafflestones. Composite Section The composite section that emerges after correlating and integrating the successions and localities mentioned above reveals a stratigraphically continuous record of the interval ranging from late Myachkovian to late Khamovnikian in the Andara Massif (Fig. 6). The uppermost Myachkovian-lowest Khamovnikian portion of the succession (the Protriticites Zone), is,300 m thick in the Ándara composite section, whereas this biostratigraphic interval does not exceed 20 m in the classic Carboniferous sections of the Moscow Basin (Alekseev et al., 2009; Goreva et al., 2009). This exceptional thickness of the Ándara Massif composite section reflects an extremely high rate of carbonate sedimentation, estimated on average ca.,30-40 cm/kyr. Due to the abundance of fusuline-bearing beds, it also represents an exceptional record of fusuline evolutionary trends. Journal of Foraminiferal Research fora d 14/7/15 13:21:26 268

6 FUSULINES FROM THE PICOS DE EUROPA 269 Protriticites Zone This biozone is represented in the uppermost part of the Picos de Europa Formation exposed in the Castillo del Grajal and Silla de Caballo Bajero sections and in the lower part (sequences 1 and 2) of the Las Llacerias Formation outcropping in the Mazarrasa and Picos del Jierro sections (Figs. 4 6). The fusuline assemblages of this interval are usually dominated by the joint occurrence of Protriticites (s. l.) and a representative of the Fusulininae (Quasifusulinoides, or, more frequently, Pseudotriticites). Accompanying genera are Ozawainella, Fusiella, Schubertella, and Pseudostaffella (very rare). The following paragraphs summarize the overall characteristics of the representatives of each genus in the studied sections. FIGURE 5. Correlation diagram of the studied sections and extent of the hiatuses detected between the Picos de Europa and Las Llacerias formations. CHARACTERISTICS OF THE FUSULINE ASSEMBLAGES OF THE LATE MOSCOVIAN AND EARLY KASIMOVIAN STRATA OF PICOS DE EUROPA Fusulinella Zone The lowest stratigraphic interval studied in this paper corresponds to the upper part of the Fusulinella Zone, the Fusulinella alvaradoi subzone (Fig. 6), equivalent to the Fusulinella B3 subzone of van Ginkel (1965). This biozone was recorded in the upper part of the Picos de Europa Formation in the Mazarrasa and the Morra de Lechugales sections (Figs. 4 5). Most characteristic forms in this interval are those belonging to the Fusulinella schwagerinoides species group, such as Fusulinella alvaradoi van Ginkel, F. loresae van Ginkel, and F. cf. curtissima Bogush, as well as F. cf. branoserae van Ginkel, F. ex gr. pseudobocki Lee & Chen, and F. aff. rara Schlykova (Figs. 6 7). One sample also provided a more diverse assemblage containing typical late Moscovian forms such as Beedeina cf. paradistenta Safonova, Fusulinella cf. valida Bogush, and Pseudostaffella (Neostaffella) ex gr. sphaeroidea von Möller, among others. The age of the Fusulinella alvaradoi subzone is late Myachkovian, but probably not the latest since this subzone is below the first appearance of the genus Protriticites. Protriticites Following van Ginkel (1971) and van Ginkel & Villa (1999), Protriticites is used here in a broad sense that includes the genera Obsoletes Kireeva, 1950, and Praeobsoletes Remizova, As discussed by van Ginkel & Villa (1999), the lower Kasimovian assemblages of the Cantabrian Zone include forms that resemble Obsoletes Kireeva but do not show the two-layered porous wall in the outer volutions that is typical for the latter genus, but rather a very thin lower tectorium is usually present. Apparently, forms exhibiting a wall microstructure similar to typical Obsoletes from the Donets Basin do not exist in the Cantabrian Zone. However, in other respects (subcylindrical shape, type of chomata, very thin wall), some forms occur that do not differ significantly from Obsoletes (e.g., Protriticites sp. 3, Fig ; P. cf. mirabilis, Fig ). Ueno (1991) has pointed out that the taxonomic status of these two genera is confused, especially with regard to spirothecal composition. However, this author suggests that, besides its thinner wall, Obsoletes can be distinguished from Protriticites in having a slightly more elongate shell and less developed chomata. These characteristics, although present in some lineages, cannot be used as reliable criteria among the Cantabrian Zone materials because some forms have Protriticites-like massive chomata (at least in the inner volutions) in Obsoleteslike elongate shells with thin walls (e.g., the two species indicated above). Praeobsoletes is not clearly established within the Cantabrian Zone fusulines. This genus was introduced to include transitional forms from Fusulinella to Obsoletes, characterized by a thin wall pierced by weak pores and consisting of tectum, dark diaphanotheca, and a thin and discontinuous lower tectorium. As the type species of this genus, Remizova (1992) designated Obsoletes burkemensis Volozhanina, In the Picos de Europa strata, this kind of wall occurs in the lower part of the Protriticites Zone, in forms here assigned to primitive Protriticites (e.g., Fig ), but it can be present in specimens occurring throughout the biozone. Moreover, one can observe that within a single species, specimens show either a Praeobsoletes or a Protriticites type of wall (the latter exhibiting a thicker lower tectorium), which show merely intraspecific variability. Baranova (2005) reexamined the type material of Obsoletes burkemensis, and other Volozhanina s (1962) Journal of Foraminiferal Research fora d 14/7/15 13:21:38 269

7 270 VILLA AND OTHERS FIGURE 6. Distribution of the fusulinid species described in this paper in the composite section resulting from the integration of the sections studied. (Correlation of the samples listed on the right is approximate). Journal of Foraminiferal Research fora d 14/7/15 13:21:44 270

8 FUSULINES FROM THE PICOS DE EUROPA 271 FIGURE 7. Fusulinella species from the Fusulinella Zone (F. alvaradoi subzone) in the uppert part of the Picos de Europa Fm. 1-4, 6-7 Fusulinella loresae van Ginkel, 1965; 1 MZ-5/1; 2 MZ-1/5; 3 MZ-1/4a; 4 MZ-6/3. 5, Fusulinella cf. curtissima Bogush, 1963, MZ-1/4b. 6 MZ-1/2; 7 MZ-1/1a. 8-9 Fusulinella ex gr. pseudobocki Lee & Chen, 1930; 8 MZ-2/4; 9 MZ-2/5. 10 Fusulinella aff. loresae van Ginkel, 1965, ML-2/ Fusulinella cf. branoserae van Ginkel, 1965; 11 MZ-2/2; 12 MZ-2/6. 13, 17 Fusulinella alvaradoi van Ginkel, 1965; 13 MZ-7/1. 14 Fusulinella aff. rara Shlykova, 1948, ML-2/5a Fusulinella ex gr. schwagerinoides Deprat, 1913; 15 ML-2/7; 16 ML-2/2; 17 MZ-2/3. species included by Remizova in Praeobsoletes (Remizova, 1992, 1993), and concluded that they form a mixed group consisting of members of Fusulinella and Protriticites s. l. On the other hand, she also pointed out the existence of morphological variability within single species, in individuals that were sometimes assigned to different genera (Baranova, 2005). In our opinion, in the transition from Fusulinella to Protriticites, an array of unstable features appeared within a single variable plexus. During Protriticites evolution, some of the more primitive characters remain in specimens from which, as stated by Remizova (1992), species showing an Obsoletes type of wall could have derived. Apparently, as mentioned above, such typical Obsoletes are not present in the Cantabrian Zone assemblages. Wahlman et al. (1997) pointed out that the same taxonomic difficulties, which have been observed in the Cantabrian Zone to separate Protriticites, Obsoletes and Praeobsoletes, and even Fusulinella, also exist with the protriticitids faunas from the western U.S.A. They noted morphotypes can be segregated, but often test and chomata shape, and wall structures, appear to vary along a continuum between end member morphotypes (p. 163). In their opinion, the frequent existence of gradational intermediate morphological features may demonstrate a high degree of genetic flexibility in the early development of the protriticitid lineage (Wahlman et al., 1997). Overall, the Protriticites species from the Ándara Massif sections here analyzed (southern part of the Picos de Europa region) are dominated by forms exhibiting a subcylindrical or an ellipsoidal outer shape. In other words, they have shells with a flat or nearly flat median region. In contrast, contemporaneous Protriticites species from the Las Llacerias section (northern part of the region, Fig. 1) are dominated by subrhomboidal and inflated forms (van Ginkel & Villa, 1999). This difference suggests that the Las Llacerias Protriticites species belong to different lineages from those of Ándara. In addition, the latter are more similar to the Protriticites studied by van Ginkel (1965) from La Pernía, an area situated southeast of the Picos de Europa region, in a different structural unit but in the same foreland basin. Despite the La Pernía and Picos de Europa samples having been taken in different carbonate systems, the similarities in the Protriticites species point to a paleogeographic connection between these areas. Pseudotriticites Protriticites is frequently associated with species of the Fusulininae, which often belong to Pseudotriticites (Fig. 10), a genus that is common in the Picos de Europa area, but appears to be rare in contemporaneous strata in other parts of the world. As noted by van Ginkel (1971), Pseudotriticites includes Beedeina-derived species characterized by a tendency to rhombic shape of the inner whorls and a regular pattern of septal folding. However, due to certain morphological convergence among Pseudotriticites and the Fusulina-descendant genus Quasifusulinoides, whose wall microstructure can be similar, it is sometimes difficult to separate the two genera. Quasifusulinoides As mentioned above, the generic assignment of many specimens placed here in Quasifusulinoides is difficult. We used several morphological criteria to distinguish that genus from Pseudotriticites. Quasifusulinoides usually exhibits a more irregular pattern of septal folding and a flatter median region of the whorls. In addition, there are also specimens that are intermediate between Quasifusulinoides and Quasifusulina. In fact, as most of the Quasifusulinoides Journal of Foraminiferal Research fora d 14/7/15 13:22:12 271

9 272 VILLA AND OTHERS FIGURE 8. Protriticites species from the lower part of the Protriticites Zone in the uppermost part of the Picos de Europa Fm. 1-3 Protriticites sp. 1; 1 CG-10/1; 2 CG-10/4; 3 CG-1/1a. 4, 6 Protriticites sp. 2 ex gr. P. grosdilovae (Miklukho Maklay, 1949). 4 CG-12/7; 6, CG-12/4. 5, 7-15 Protriticites grajalensis Villa sp. nov. 5 CG-15/8; 7 CG-14/8a; 8 CG-15/7; 9 CG-15/5; 10 CG-15/2a; 11 CG-15/6b; 12 CG-15/10d; 13 CG-15/4, holotype; 14 CG-15/ 6a; 15 CG-15/3b Protriticites sp. 3 ex gr. P. grosdilovae (Miklukho-Maklay, 1949); 16 CG-22/9a; 17 CG-20/2; 18 CG-21/4; 19 CG-21/2; 20 CG- 22/5; 21 CG-14/5a Protriticites quasiovoidalis Villa sp. nov; 22 CG-22/6, holotype; 23 CG-22/2; 24 CG-CG-22/4b; 25 CG-22/1; 26, CG-24/ Protriticites ex gr. subschwagerinoides Rozovskaja, 1950; 27 SCC-1/4; 28 SCC-1/ Protriticites aff. arsenjevi (Nikitina, 1969); 29 CG-23/4; 30 CG-23/6; 31 CG-23/ Protriticites cf. manukalovae Kireeva, 1950; 32 CG-12/1;33 CG-16/2. 34 Protriticites aff. vetus (Rjazanov, 1958), SCB-1/1. species found in the Ándara Massif show a very thin wall (Fig. 11), sometimes with almost negligible tectoria, their assignment to Quasifusulina is conceivable. This is particularly the case in forms from the Protriticites Zone, which show an overall resemblance to Quasifusulinoides but possess thin two-layered walls and a large proloculus. Those features suggest a closer affinity to Quasifusulina, but this genus is usually recorded in late Kasimovian and Gzhelian strata. Such morphotypes, here questionably assigned to Quasifusulinoides (Fig. 11), could be Journal of Foraminiferal Research fora d 14/7/15 13:22:14 272

10 FUSULINES FROM THE PICOS DE EUROPA 273 FIGURE 9. Protriticites species from the upper part of the Protriticites Zone in the lower part of the Las Llacerias Fm Protriticites benshae Villa sp. nov. 1 MZ-9/2; 2 MZ-9/10b; 3 MZ-9/8; 4 MZ-9/10a; 5 MZ-9/9; 6 MZ-9/5, holotype; 7 MZ-9/4; 8 MZ-9/6; 9 AND-8/4b; 10 AND-8/ Protriticites aff. confusus (Kireeva, 1950). 11 PJ-0/7; 12 PJ-0/1; 13 PJ-0/8. 14 Protriticites cf. subschwagerinoides Rozovskaja, 1950, PJ-0/13a Protriticites cf. mirabilis (Kireeva, 1950). 15 PJ-1/7; 16 PJ-1/2b; 17 PJ-1/6a Protriticites? sp. 4; 18 CG-29/7; 19 CG-29/1; 20 CG-29/5; 21 CG-29/ 4; 22 CG-29/2. 23 Protriticites winkleri Villa, 2010, PJ-5/1 (specimen LE5-F5/9 in Martínez-Chacón et al., 2010). 24 Protriticites? sp. 5, CG-31/3. alternatively considered either ancestors of Quasifusulina or primitive representatives of this genus. Other Genera Although not described in this paper, the occurrence of other fusuline genera must be noted. The lower part of the Protriticites Zone (samples CG-12 and CG-21, respectively; Fig. 6) has yielded a few small specimens of Pseudostaffella exhibiting only 4.5 and 3.5 volutions. These small-sized Pseudostaffella are not juvenaria, for they occur constantly in the latest Maychkovian and early Kreviakinian strata (see van Ginkel & Villa, 1991 for Kreviakinian specimens from the Picos del Jierro section). As many Russian authors restricted the genus Pseudostaffella for the Bashkirian and early Moscovian forms, and included the larger, late Moscovian species in the genus Neostaffella, Solovieva (1986) created the genus Quasistaffella for smaller species occurring in the Krevyakinian. In our opinion, Quasistaffella is morphologically indistinguishable from Pseudostaffella. On the other hand, intermediate forms between typical Pseudostaffella and typical Neostaffella are abundant. For these reasons, we think it would be advisable to preserve the whole group within a single genus, Pseudostaffella. Nevertheless, as within the range of variability one finds that the extreme morphotypes of Pseudostaffella and Neostaffella are really different, we tentatively use all these names as subgenera. Fusiella are rarely found through the whole Protriticites Zone. Unfortunately, most of them are oblique sections that make species identification difficult. The exceptions are two broken specimens from locality MZ-10 of the Mazarrasa section, which are very close to or belong to F. lancetiformis Putrja, and a few oblique specimens of a Fusiella species resembling F. pseudorawi Isakova. According to the distribution and phylogenetical Journal of Foraminiferal Research fora d 14/7/15 13:22:18 273

11 274 VILLA AND OTHERS FIGURE 10. Beedeina and Pseudotriticites species from the Picos de Europa (upper part) and Las Llacerias formations. 1 Beedeina cf. paradistenta (Safonova, 1951), MZ-4/1. 2 Pseudotriticites? sp. 1, CG-16/4. 3-6, 8, 11 Pseudotriticites cantabricus Villa sp. nov. 3 CG-24/3; 4 CG-16/3; 5 CG-14/2; 6 CG-19/3; 8 CG-19/4a; 11 CG-19/5. 7, 9-10, Pseudotriticites aff. oblonga (Nikitina, 1969). 7 PJ-0/17; 9 PJ-0/11; 10 MZ-10/2a; 12 PJ-0/15; 13 PJ-0/ 5; 14 PJ-0/3. 15 Pseudotriticites sp. 2, AND-3/3b. relationships of the Fusiella species proposed by Isakova (2013), the coexistence of F. lancetiformis and F. pseudorawi in MZ-10 would confirm the late Kreviakinian age of these beds. Schubertella fragments, mainly from the S. kingi and S. obscura groups, as well as diverse Ozawainella specimens (O. rhomboidalis and O. pseudoangulata groups), also occur at several levels through the Protriticites Zone. Montiparus Zone Fusulines belonging to the lower part of the Montiparus Zone occur at the Castillo del Grajal section, in strata of the Las Llacerias Formation resting unconformably on Protriticites beds of the Picos de Europa Formation. They are isolated specimens of Montiparus ex gr. umbonoplicatus (Fig. 12.5) as well as a species transitional between Protriticites and Montiparus that has been tentatively identified as Protriticites? sp. 4 (Fig ) and Protriticites? sp. 5 (Fig. 9.24). More diverse and advanced Montiparus species occur at other sites of the Ándara Massif exposing the upper part of the Las Llacerias Formation (Silla de Caballo Cimero section and AND- localities in the Grajales Ridge; Figs. 3, 4). There we found Montiparus belonging to the Journal of Foraminiferal Research fora d 14/7/15 13:22:22 274

12 FUSULINES FROM THE PICOS DE EUROPA 275 FIGURE 11. Quasifusulinoides and Quasifusulina species from the Picos de Europa (upper part) and Las Llacerias formations. 1-4, 6 Quasifusulinoides? sp CG-14/2b; 2 CG-14/6d; 3 CG-14/8b; 4 CG-16/1; 6 CG-15/11a. 5 Quasifusulinoides sp. 2, CG-15/3a. 7-9, 13 Quasifusulinoides ex gr. parafusiformis Bensh, 1969; 7 SCB-2/1; 8 SCB-2/2; 9 SCB-3/2; 13 SCB-2/ Quasifusulinoides parafusiformis Bensh, 1969; 10 CG-27/4; 11 CG-27/3. 12 Quasifusulina praecursor Rauzer-Chernousova, 1951, AND-0/3. 14 Quasifusulina sp. 1, AND-3/2b. 15 Quasifusulina sp. 2, AND-3/4. M. umbonoplicatus and M. paramontiparus species groups, and a slender species closely related to Montiparus fischeri van Ginkel (Fig. 12). The latter exhibits an advanced type of wall (thick pores in the outer volutions), indicating an evolutionary stage close to the Montiparus-descendant genus Rauserites. These younger beds of the Grajales Ridge and Silla de Caballo Cimero peak also contain large Quasifusulina, isolated Pseudotriticites, Fusiella, and Ozawainella specimens. SYSTEMATIC PALEONTOLOGY In this section we describe 41 fusulinoidean taxa including four new species, providing measurements of the following parameters: L, length of the outer shell; D, Journal of Foraminiferal Research fora d 14/7/15 13:22:27 275

13 276 VILLA AND OTHERS FIGURE 12. Montiparus species from the Montiparus Zone in the upper part of the Las Llacerias Formation. 1, 3 Montiparus aff. umbonoplicatus (Rauzer-Chernousova & Beljaev, 1937); 1 AND-1/6; 3 AND-1/5. 2 Montiparus paramontiparus (Rozovskaja, 1950), AND-1/2. 4, 7-8 Montiparus cf. umbonoplicatus (Rauzer-Chernousova & Beljaev, 1937); 4 SCC-2/1; 7 SCC-2/6; 8 SCC-2/2. 5 Montiparus ex gr. umbonoplicatus (Rauzer-Chernousova & Beljaev, 1937), CG-28/3a. 6, 9-11 Montiparus aff. fischeri van Ginkel, 1971; 6 AND-2/2; 9 AND-2/1; 10 SCC-3/2b; 11 SCC-3/2a. diameter of the outer shell; L/D, length/diameter ratio; n, number of volutions; D IV, diameter of the fourth whorl; d, diameter of the proloculus; wth, wall thickness referred to the penultimate volution. Figured specimens are deposited in the micropaleontological collection of the Departamento de Geología, Universidad de Oviedo (Spain). Order FORAMINIFERIDA von Eichwald, 1830 Suborder FUSULININA Wedekind, 1937 Superfamily FUSULINOIDEA von Möller, 1878 Family FUSULINIDAE von Möller, 1878 Subfamily FUSULINELLINAE von Staff & Wedekind, 1910 Genus Fusulinella von Möller, 1877 Type species: Fusulinella bocki von Möller, 1877 Fusulinella loresae van Ginkel, 1965 Fig , Fusulinella loresae van Ginkel, 1965, p , pl. 41, figs , pl. 42, figs Measurements. L mm; D mm; L/D ; n56 6.5; D IV mm; d mm; wth mm. Remarks. Test shape varies from ovoidal to fusiform. Spire tightly coiled in the first 3.5 whorls, and rapidly expanding in the succeeding ones. Axis of the first volutions arranged at a large angle to outer volutions. Chomata irregular, asymmetrical, frequently low and narrow, sometimes broader and more prominent. Septa undulated in their polar ends. Wall consisting of four layers, including a very thin diaphanotheca and highly variable tectoria. In the outer volutions the lower tectorium may be as thick as the rest of the layers. Fusulinella loresae, originally described from La Pernía area (northern Palencia, Cantabrian Zone), resembles some of the species included by Bogush (1963) in the F. schwagerinoides species group, especially the specimens assigned by this author to F. adjuncta Schlykova, However, F. loresae can be distinguished by its more tightly coiled juvenarium and more expanded outer volutions. This species also resembles F. alvaradoi van Ginkel, from which F. loresae differs in having a smaller size on average, and less expanded outer volutions. It is also reminiscent of F. hanzawai Igo, 1957, from which it can be distinguished by its stronger septal fluting. Stratigraphic data and age. Samples MZ-1, MZ-5 and MZ-6, Picos de Europa Formation, Mazarrasa section. Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Fusulinella aff. loresae van Ginkel, 1965 Fig Measurements. L54.15 mm; D51.45 mm; L/D52.86; n55.5; D IV mm; d570 mm; wth535 mm. Remarks. The fast spiral growth indicates the affinity of this specimen with the Fusulinella schwagerinoides species group. In its size, elongated fusiform shape and thin spirotheca, it very much resembles F. loresae. However, this material differs from van Ginkel s (1965) types in having a somewhat more subrhombic shape of the outer whorls and less tight juvenarium. Stratigraphic data and age. Sample ML-2, Picos de Europa Formation, Morra de Lechugales section. Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Journal of Foraminiferal Research fora d 14/7/15 13:22:32 276

14 FUSULINES FROM THE PICOS DE EUROPA 277 FIGURE 13. Spirotheca of Fusulinella, Protriticites and Montiparus species from the Ándara Massif sections. 1 Fusulinella alvaradoi van Ginkel, 1965, MZ-7/1 (in Fig. 7.13). 2-3 Protriticites sp CG-1/1a (in Fig. 8.3); 3 CG-10/1 (in Fig. 8.1). 4 Protriticites cf. mirabilis (Kireeva, 1950), PJ-1/7 (in Fig. 9.15). 5 Protriticites benshae Villa sp. nov, MZ-9/5 (in Fig. 9.6). 6 Protriticites? sp. 4, CG-29/5 (in Fig. 9.20). 7 Montiparus aff. umbonoplicatus (Rauzer-Chernousova & Beljaev, 1937), AND-1/ Montiparus aff. fischeri van Ginkel, 1971; 8 AND-2/1 (in Fig. 12.9); 9 SCC-2/2. Fusulinella cf. curtissima Bogush, 1963 Fig. 7.5 Measurements. L52.00 mm; D51.00 mm; L/D52; n55.5(?); D IV mm. Remarks. Juvenarium and spiral coiling of this single specimen resemble those of F. loresae (present in the same sample), from which it differs in having smaller size, smaller L/D ratio and smaller chomata. All these characters, as well as the shape of the shell and type juvenarium are similar to those of F. curtissima. Stratigraphic data and age. Sample MZ-1, Picos de Europa Formation, Mazarrasa section. Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Fusulinella alvaradoi van Ginkel, 1965 Figs. 7.13, 7.17, 13.1 Fusulinella schwagerinoides alvaradoi van Ginkel, 1965, p , pl. 42, figs , pl. 43, figs Measurements. L mm; D mm; L/D ; n55.5 6; D IV mm; d mm; wth mm. Remarks. This species, described originally from the Cantabrian Mountains by van Ginkel (1965) as F. schwagerinoides alvaradoi, can easily be identified by its tiny proloculous and tightly coiled juvenarium, and, especially, the high rate of spiral growth. Chomata varies from asymmetric, narrow and relatively high (Fig. 7.17) to low and rounded (Fig. 7.13); such a variability is also observed in the material by van Ginkel (1965). Tunnel opening very wide. Wall usually thin and consisting of four layers, of which a remarkable feature is the very thin but distinct diaphanotheca (Fig. 13.1). A very similar species is F. altispiralis Bogush, 1963, from which F. alvaradoi apparently only differs in having somewhat stronger chomata. Stratigraphic data and age. Samples MZ-2, MZ-7, Picos de Europa Formation, Mazarrasa section. Fusulinella Zone, Journal of Foraminiferal Research fora d 14/7/15 13:22:34 277

15 278 VILLA AND OTHERS Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Fusulinella ex gr. schwagerinoides Deprat, 1913 Figs Measurements. L mm; D mm; L/D ; n54.5? 5.5; D IV mm; d mm; wth540 mm. Remarks. Our material has a tightly coiled juvenarium followed by rapidly expanding volutions and narrow chomata that are typical for the Fusulinella schwagerinoides species group. The present specimens are very close to those of Fusulinella alvaradoi, but differ mainly in having smaller shells. Stratigraphic data and age. Samples ML-2, Picos de Europa Formation, Morra de Lechugales section. Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Fusulinella ex gr. pseudobocki Lee & Chen in Lee et al., 1930 Figs Measurements. L mm; D mm; L/ D ; n55.5 6; D IV mm; wth mm. Remarks. Shell oval to short-fusiform, with blunt-pointed polar ends, tiny proloculus and tightly coiled inner volutions. Chomata broad in the inner whorls, becoming higher and of quadrangular shape in the outer ones. Wall consisting of four layers: tectum, diaphanotheca and fairly thick tectoria. This species resembles forms of the F. pseudobocki and F. mosquensis groups (e. g. F. ovoides Rauzer-Chernousova in Rauzer-Chernousova et al., 1951, and F. maldrigensis van Ginkel, 1965), but differs from all of them by its smaller size. Stratigraphic data and age. Sample MZ-2, Picos de Europa Formation, Mazarrasa section. Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Fusulinella cf. branoserae van Ginkel, 1957 Figs Measurements. L mm; D mm; L/ D53.6; n56.5; D IV mm; d mm; wth mm. Remarks. Studied material consists only of two specimens sharing an elongated subcylindrical test, inner 3.5 whorls tightly coiled and with first 1.5 whorls coiled at a large angle with respect to subsequent ones, and wall consisting of four layers. However, they differ from one another in the development of epithecal deposits, for chomata and tectoria are better developed in the specimen MZ-2/6 (Fig. 7.12). In this respect, it is remarkable that the last two volutions of the other specimen (MZ-2/2, Fig. 7.11) show a very thin and dark undifferentiated wall that is only 20 mm thick. Both specimens are tentatively placed into a single species and compared to F. branoserae, to which they show the greatest resemblance (especially specimen MZ-2/6, Fig. 7.12). Somewhat similar species are also F. gracilis Kanmera, 1954, and F. mira Semikhatova, Stratigraphic data and age. Sample MZ-2, Picos de Europa Formation, Mazarrasa section. Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Fusulinella aff. rara Schlykova, 1948 Fig Measurements. L55.60 mm; D52.00 mm; L/D52.80; n56; D IV mm; d590 mm; wth540 mm. Remarks. For the large L/D ratio, subcylindrical shape, flat median region, and type of chomata, this single specimen resembles forms belonging to the Fusulinella mosquensis species group, especially F. rara. Noteworthy is the resemblance to the materials from the Alay Range assigned by Bogush (1963) to this species, from which the Picos de Europa specimen differs only in having a slightly larger size for corresponding whorls. Stratigraphic data and age. Sample ML-2, Picos de Europa Formation, Morra de Lechugales section. Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Genus Protriticites Putrja, 1948 Type species: Protriticites globulus Putrja, 1948 Protriticites sp. 1 Figs , Measurements. L ? mm; D mm; L/D ; n55.5 7; D IV mm; d mm; wth mm. Remarks. Shape of the shell from ellipsoidal to subcylindrical. Chomata high (reaching at least half the chamber height) and ribbon-shaped, extending to the poles in the inner whorls; in the outer whorls, the chomata are narrower and more irregular. Wall transitional from Fusulinella to Protriticites, pierced by weak pores (sometimes undetectable) and consisting of four layers: tectum, gray diaphanotheca, and very thin upper and lower tectoria (Fig. 13.2). Most distinct features of this species are its wall microstructure (rather primitive for the genus Protriticites), flat median region of the whorls, and well-developed chomata. The characteristics of the wall are similar to those of the forms included by Remizova (1992) in the genus Praeobsoletes. The specimens described here somewhat resemble the types of P. lutugini Kireeva, 1949, and P. nikitovkensis Kireeva, 1949, but differ in having a flatter median region and probably a more primitive wall. Stratigraphic data and age. Samples CG-1, CG-2, CG-4, CG-6, CG-10, CG-11, CG-12, Picos de Europa Formation, Castillo del Grajal section. Lowermost part of the Protriticites Zone, Moscovian/Kasimovian transition interval, either top of Myachkovian or lowermost Krevyakinian. Protriticites sp. 2 ex gr. P. grosdilovae (Miklukho-Maklay, 1949) Figs. 8.4, 8.6 Measurements. Specimen CG-12/4 (Fig. 8.6): L55.45 mm; D51.67 mm; L/D:53.26; D IV mm; d580 mm; wth540 mm. Remarks. This species is reminiscent of Protriticites mirabilis (Kireeva, 1950) and P. grosdilovae in having a shell Journal of Foraminiferal Research fora d 14/7/15 13:22:37 278

16 FUSULINES FROM THE PICOS DE EUROPA 279 exhibiting a flat median region. However, the microstructure of the wall is more primitive than in the abovementioned species, being closer to Fusulinella and to transitional species assigned to Praeobsoletes by Remizova (1992). Furthermore, Protriticites sp. 2 is less elongated and exhibits more regular volutions than P. mirabilis and has weaker septal folding than P. grosdilovae. Somewhat similar species are Protriticites sp. 1 and Protriticites grajalensis sp. nov., both described in this paper from the Castillo del Grajal section. Protriticites sp. 2 differs from the former in having less massive (narrower and lower) chomata, and from the latter in exhibiting a flatter median region of the whorls. Stratigraphic data and age. Sample CG-12, Picos de Europa Formation, Castillo del Grajal section. Lower part of the Protriticites Zone, Moscovian/Kasimovian transition interval (either top of Myachkovian or lowermost Krevyakinian). Protriticites sp. 3 ex gr. P. grosdilovae (Miklukho-Maklay, 1949) Figs Measurements. L mm; D mm; L/D ; n ; D IV mm; d mm; wth mm. Remarks. The main characteristics of the available specimens are their subcylindrical shell and fast growth of the spire in the last 1.5 whorls, resulting in a diameter of the fifth whorl that nearly doubles that of the fourth. Chomata high and broad in the inner volutions, narrow and low in the last ones. Wall very thin, pierced by fine pores, and consisting of tectum, diaphanotheca, very thin inner tectorium (sometimes virtually indiscernible), and a discontinuous and thin outer tectorium that is sometimes only present in the inner volutions. This species differs from all known forms by the combination of loose-coiled outer whorls, massive chomata in the inner volutions and a thin wall. However, the scarcity of well-preserved specimens prevents us from describing it formally as a new species. For its subcylindrical adult shape, high spire of outer whorls and rounded and broad polar ends, Protriticites sp. 3 is close to P. grosdilovae, of special note being the resemblance between the specimen illustrated in Fig and the Central Asian specimen of P. grosdilovae illustrated by Bensh (1972; pl. III, fig. 6). However, the Picos de Europa form differs from the the types of P. grosdilovae by Miklukho-Maklay (1949) in having a larger proloculus, less plicated septa, a non-depressed central part, and perhaps a more primitive wall microstructure (Fig. 13.3). Less similar species are Protriticites tadushensis (Nikitina, 1969) and P. kireevae (Grozdilova, 1966). Stratigraphic data and age. Samples CG-14, CG-20, CG- 21, CG-22, Picos de Europa Formation, Castillo del Grajal section. Protriticites Zone, lower Kasimovian (Krevyakinian). Protriticites grajalensis Villa sp. nov. Figs. 8.5, Material. 4 axial, 1 nearly axial, 1 paraxial, 1 slightly oblique, and a number of oblique sections. Holotype: specimen CG-15/4 (Fig. 8.13). Derivatio nominis: From the name of the Castillo del Grajal peak. Measurements. L mm; D mm; L/D ; n55.5 7; D IV mm; d mm; wth mm. Holotype: L56.20 mm; D52.35 mm; L/D52.64; n57; D IV mm; d5100 mm; wth540 mm. Description. Test of rather large size and variable outer shape; most of the specimens show a flat middle region in the last volution and, therefore, the outer shell is subcylindrical, whereas in others the last volution is slightly convex and so the test is elongated fusiform. In contrast, the median region of the inner and intermediate volutions are convex in all the specimens. Chomata broad and of variable shape, frequently subquadrangular, their height usually being half of the chamber height. Septal folding moderate, developed only along the axial region. Wall consisting of four layers, with very thin tectoria in the inner and intermediate whorls, and three layers in the outer ones, where the outer tectorium is absent. Discussion. Although the study of the present specimens is hampered by the poor state of preservation of the outer whorls, the characteristics of the species clearly differ from those of the described protriticitids. For its large size and tendency to develop a subcylindrical shell, the species resembles Protriticites mirabilis, P. lamellosus Kireeva, 1950, and P. grosdilovae, but it differs from them in having broader chomata, less intense septal folding, and larger proloculus. Besides, P. mirabilis has tighter coiling in the inner whorls and more rapid expansion of the spire in the outer ones. Type of chomata and shape of the inner volutions also resemble P. manukalovae Kireeva, 1950, but the latter has a shorter shell and smaller proloculus. Some specimens (e.g., Figs and 8.14) are somewhat reminiscent of Obsoletes? formosus Progryebnyak, 1975, which differs in having a smaller shell, tighter spire and probably a more advanced wall microstructure. From Protriticites sp. 2 described earlier, P. grajalensis sp. nov. can be distinguished by having a convex median region of the whorls and a looser spiral in the outer volutions. Stratigraphic data and age. Samples CG-14, CG-15, Picos de Europa Formation, Castillo del Grajal section. Lower part of the Protriticites Zone, Moscovian/Kasimovian transition interval or lowermost Krevyakinian. Protriticites quasiovoidalis Villa sp. nov. Fig Material. 7 axial, 1 paraxial, 2 slightly oblique, and a number of oblique sections. Holotype: specimen CG-22/6 (Fig. 8.22). Derivatio nominis. Making reference to its ovoid shape in section. Measurements. L mm; D mm; L/D ; n ; D IV mm; d mm; wth mm. Holotype: L55.10 mm; D52.10 mm; L/D52.43; n56.75; D IV mm; d560 mm; wth570 mm. Description. Shell short-fusiform, short planar-fusiform or short subcylindrical, frequently exhibiting an ovoid shape in section. Proloculus small, first one to two volutions tightly coiled, then expanding uniformly. Chomata well-developed, frequently asymmetrical and of variable shape, from broad Journal of Foraminiferal Research fora d 14/7/15 13:22:37 279

17 280 VILLA AND OTHERS and subquadrangular, to narrow and rounded or wedgeshaped. Septa folded in the polar regions; exceptionally, some loops may also occur in the lateral sides. Wall quite thick in the genus and pierced by thin pores, consisting of tectum, gray diaphanotheca and irregular and discontinuous tectoria. Discussion. The present new species resembles P. burkemensis (Volozhanina, 1962) and P. spectabilis (Volozhanina, 1962). It differs from the former in having a smaller proloculus and smaller diameter in the inner volutions, slightly larger shell size in mature specimens, and thicker spirotheca. From P. spectabilis, the Picos de Europa species is distinguished by its well-developed chomata, less intense septal fluting, and thinner wall. Another similar form is Protriticites subcylindricus (Bogush, 1960), from which P. quasiovoidalis can be separated by its smaller L/D ratio and fewer volutions. The inner stage of P. quasiovoidalis sp. nov. is reminiscent of the specimens of Protriticites sp. 3 described above. However, the latter clearly differs in exhibiting a much faster spiral growth in the last volutions. Stratigraphic data and age. Samples CG-22, CG-24, CG- 25, Picos de Europa Formation, Castillo del Grajal section. Protriticites Zone, lower Kasimovian (Krevyakinian). Protriticites ex gr. subschwagerinoides Rozovskaja, 1950 Fig Measurements. L, mm; D mm; L/D, ; n55 7; D IV mm; d mm; wth mm. Due to poor preservation or orientation of available specimens, measurements are approximate. Remarks. Fusiform to elongate fusiform shell, exhibiting tightly coiled inner volutions and looser outer ones. Specimen illustrated in Fig has the first volution coiled at an angle to subsequent ones. Septa folded in their axial ends. Chomata usually narrow and subquadratic, rather irregular and of moderate height. Wall consisting of four layers in intermediate whorls; in the outer volutions, the outer tectorium is reduced or absent. The type of coiling and the type of chomata point to the close affinity of this species to forms allied to the P. subschwagerinoides group, to which somewhat similar species are Protriticites variabilis Bensh, 1972, and P. subovatus Bensh, 1972, both from Central Asia. Also similar is P. subcylindricus (Bogush, 1960), although the latter has a flatter median region. Stratigraphic data and age. Sample SCC-1, top of the Picos de Europa Formation, Silla de Caballo Cimero section. Protriticites Zone, Kasimovian (Krevyakinian). Protriticites aff. arsenjevi (Nikitina, 1969) Fig Measurements. L mm; D mm; L/D ; n54.5 5; D IV mm; d mm; wth mm. Remarks. Size and shape of shell, septal fluting and type of coling of this species are similar to characteristics of P. arsenjevi. However, it differs in having a distinct inner tectorium, which is discontinuous in P. arsenjevi, and lower chomata. Stratigraphic data and age. Sample CG-23, Picos de Europa Formation, Castillo del Grajal section. Protriticites Zone, Kasimovian (Krevyakinian). Protriticites cf. manukalovae Kireeva, 1950 Figs Measurements. L mm; D mm; L/D ; n56.5 8; D IV mm; d560 mm; wth mm. Remarks. The present specimens closely fit the characteristics of Protriticites manukalovae, particularly with regard to the ovoid shape of the shell, minute proloculus, relatively strong septal fluting in the axial ends of septa, massive chomata, and primitive microstructure of the wall, which seems to be transitional between Fusulinella and Protriticites. They only differ in having a larger number of volutions (6.5 8 whorls, opposed to 5.5 in the types of Kireeva, 1950), and, consequently, a larger shell size. Moreover, the Picos de Europa specimens have thinner tectoria, which are thinner than the diaphanotheca. Stratigraphic data and age. Samples CG-12, CG-16, Picos de Europa Formation, Castillo del Grajal section. Lower part of the Protriticites Zone, Moscovian/Kasimovian transition interval (either top of Myachkovian or lowermost Krevyakinian). Protriticites aff. vetus (Rjazanov, 1958) Fig Measurements. L55.65 mm; D52.25 mm; L/D52.51; n56?; D IV mm; wth535 mm. Remarks. Specimen of paraxial section showing a short subcylindrical shell, tiny proloculus, subquadrangular chomata (narrow and of moderate height), and possibly phrenothecae. Very thin wall, consisting of two layers irregularly underlined by a discontinuous inner tectorium; it does not show porosity. The specimen reminds us of P. vetus, particularly the holotype by Rjazanov (1958) illustrated in pl. 1, fig. 18 because of the shape of the shell, L/D ratio, and perhaps the wall microstructure, but differs in exhibiting stronger chomata and strongly folded septa at the polar ends. Protriticites winkleri Villa, in Martínez-Chacón et al., 2010, present in higher strata of the Las Llacerias Formation, is also somewhat similar; however, the latter differs in having a tighter spiral, slightly smaller size, weaker chomata, and more advanced wall (represented by distinct porosity). For the shape of the shell, type of coiling and presence of phrenothecae, it is also reminiscent of the specimens from the Carnic Alps assigned by Forke & Samankassou (2000) to Protriticites? inflatus Bensh, 1972, of which, however, differ by having more massive chomata. Stratigraphic data and age. Sample SCB-1, top of the Picos de Europa Formation, Silla de Caballo Bajero section. Protriticites Zone, Kasimovian (Krevyakinian). Protriticites benshae Villa sp. nov Fig , 13.5 Protriticites lamellosus Kireeva: Bensh, 1972, p , pl. 2, figs Journal of Foraminiferal Research fora d 14/7/15 13:22:37 280

18 FUSULINES FROM THE PICOS DE EUROPA 281 Material. 5 axial or nearly axial, 1 eccentric, 1 slightly oblique, and a number or oblique sections. Holotype: specimen MZ-9/5 (Figs. 9.6, 13.5). Derivatio nominis. The species is dedicated to Dr. F. R. Bensh. Measurements. L mm; D mm; L/D ; n55 7.5; D IV mm; d mm; wth mm. Holotype: L55.25 mm; D mm; L/D5 2.84; n5 6.5; D IV mm; d5 75 mm; wth5 65 mm. Description. Test shape rather variable, both in outer shape and L/D ratio; it is usually fusiform with polar ends varying from rounded to pointed. Chomata of quadrangular shape, usually extending to the poles in the inner twothree volutions, and narrower, sometimes more rounded, and of moderate height in the outer ones. Septal folding restricted to the axial region. Microstructure of the wall as in typical Protriticites, that is, it consists of tectum, gray diaphanotheca, and inner and outer tectoria. A remarkable feature in this species is the presence of septal lamellae (phrenothecae), which occur in all specimens. Discussion. Protriticites benshae sp. nov. resembles P. lamellosus Kireeva, 1950, from the Donets Basin in the shape of the shell, and, especially, in having phrenothecae. However, it clearly differs from the latter in having weaker septal folding. In this respect, the Picos de Europa material more closely resembles the specimens from Central Asia (Fergana) that Bensh (1972) assigned to P. lamellosus Kireeva (compare our specimen illustrated in Fig. 9.1 and that of Bensh, 1972, pl. 2, fig. 1). The Central Asian specimens are considered here to be conspecific with P. benshae sp. nov. Some of the P. benshae specimens also recall in shape, chomata and septal folding Protriticites subschwagerinoides Rozovskaja, 1950, but the latter clearly differs in lacking phrenothecae. Stratigraphic data and age. Samples MZ-9, Las Llacerias Formation, Mazarrasa section, and AND-8, Las Llacerias Formation, isolated locality in the Grajales Ridge. Upper part of the Protriticites Zone, Kasimovian (upper Krevyakinian). Protriticites aff. confusus (Kireeva, 1950) Fig Measurements. L mm; D mm; L/D ; n ; D IV mm; d mm; wth mm. Remarks. Specimens show fusiform to subcylindrical shell, with tightly coiled inner whorls, rather large L/D ratio, flat median region, and moderate to strong chomata that extend to the poles in several inner volutions. The wall is thin, four-layered, and pierced by indistinct pores; it consists of tectum, distinct diaphanotheca, and very thin tectoria, irregularly distributed but present even in the outer whorls. In their general outline, this species is similar to some elongated forms usually assigned to the genus Obsoletes, e.g., Protriticites confusus, P. gapeevi (Kireeva, 1950), and P. normalis (Nikitina, 1969). However, the Picos de Europa form differs from all of them in having, apparently, a less advanced wall microstructure and a larger shell. Stratigraphic data and age. Sample PJ-0, Las Llacerias Formation, Picos del Jierro section. Upper part of the Protriticites Zone, Kasimovian (upper Krevyakinian). Protriticites cf. subschwagerinoides Rozovskaja, 1950 Fig Measurements. L54.25 mm; D51.45 mm; L/D52.93; n55.5; D IV mm; d5100 mm; wth540 mm. Remarks. Shell subrhombic in the last two whorls and tightly coiled in all volutions except for the last one, which expands and elongates rapidly. Chomata massive and extending to the poles in the inner volutions, narrower in the penultimate whorl. Septal folding moderate and restricted to the polar ends. Wall consisting of four layers, including tectum, gray diaphanotheca and thin tectoria. This single specimen is quite similar to Protriticites subschwagerinoides, from which it only differs in exhibiting a dark but distinct diaphanotheca. It differs from the specimens from the Picos de Europa Formation assigned above to P. ex gr. subschwagerinoides in showing more acute polar ends, broader chomata in the inner volutions, and looser spire. Stratigraphic data and age. Sample PJ-0, Las Llacerias Formation, Picos del Jierro section. Upper part of the Protriticites Zone, Kasimovian (upper Krevyakinian). Protriticites cf. mirabilis (Kireeva, 1950) Fig , 13.4 Measurements. L mm; D mm; L/D ; n55 6(?); D IV (?) mm; d mm; wth mm. Remarks. Test subcylindrical, with the whorls very rapidly elongating and growing in height in each volution. Proloculus minute. Chomata irregular, often asymmetrical, narrow and high, subquadrangular in the intermediate whorls. Septa very thin and moderately folded. Phrenothecae are visible in some specimens. Wall four-layered (Fig. 13.4), usually very thin (20 30 mm) in the intermediate volutions. Tectoria irregular and frequently thin, present in all mature volutions except in the last one, which shows a two-layered wall and distinct porosity. These characteristics, including the wall microstructure, closely correspond to those of P. mirabilis (Kireeva, 1950), a species originally described from the O 1 limestone of the Donets Basin. Less similar is P. cylindricus (Kireeva, 1950), from which the Picos de Europa form differs in having stronger chomata. Stratigraphic data and age. Sample PJ-1, Las Llacerias Formation, Picos del Jierro section. Upper part of the Protriticites Zone, Kasimovian (upper Krevyakinian). Protriticites winkleri Villa in Martínez-Chacón et al., 2010 Fig Protriticites winkleri Villa in Martínez-Chacón et al., 2010, p , pl. 6, figs Measurements. L mm; D mm; L/D ; n55 5.5; D IV mm; d mm; wth mm. Remarks. The short ellipsoidal shape of the shell, with flat and parallel median region of the volutions, and the low Journal of Foraminiferal Research fora d 14/7/15 13:22:37 281

19 282 VILLA AND OTHERS to moderate irregular chomata are the most distinctive features of this species. In the mature volutions, except the last one, its wall consists of four layers: tectum, a dark diaphanotheca, and thin tectoria; thin pores pierce all layers. Stratigraphic data and age. Sample PJ-2 and PJ-5 (5LE5- F5 in Martínez-Chacón et al., 2010), Las Llacerias Formation, Picos del Jierro section. Upper part of the Protriticites Zone, Kasimovian (uppermost Krevyakinianlowermost Khamovnikian). Protriticites? sp. 4 Figs , 13.6 Measurements. L mm; D mm; L/D ; n55 6.5; D IV mm; d mm; wth mm. Remarks. Test elongate-fusiform to subcylindrical. Chomata varying from broad and rather low to narrow and high. Wall pierced by fine pores, and consisting of tectum, primatheca and outer tectorium in two inner whorls; in the subsequent volutions the outer tectorium reduced, nearly disappearing, and the primatheca is replaced by a gray diaphanotheca underlined by a thin inner tectorium (Fig. 13.6). Although Picos de Europa material conforms well to the shape and size of the shell, L/D ratio, and type of chomata of Protriticites confusus, it differs in having an inner tectorium, as well as a wall porosity perhaps weaker than those from the Donets Basin. Judging from its spirothecal microstructure, this species seems to be transitional from Protriticites to Montiparus. Stratigraphic data and age. Sample CG-29, Las Llacerias Formation, Castillo del Grajal section. Montiparus Zone, Kasimovian (lower Khamovnikian). Protriticites? sp. 5 Fig Measurements. L.5.25 mm; D51.80 mm; L/D.2.92; n56.5; D IV mm; d5110mm; wth560mm. Remarks. Test ellipsoidal to subcylindrical, tightly coiled,withflatmedianregionintheintermediateand outer whorls. Chomata rather massive, high and broad, extending to the poles in the first four volutions, then narrowing. Septa fluted in their axial region. Wall pierced by fine pores and consisting of three layers (outer tectorium, tectum and primatheca) in the 3.5 inner volutions, four layers by adding an inner tectorium below the primatheca/diaphanotheca in the intermediate whorls, and two layers in the last whorl. Protriticites? sp. 5 differs from all known Protriticites and Montiparus species in the combination of these features. The transitional character between Protriticites and Montiparus is reinforced by the presence of a likely conspecific specimen found in the same sample, which shows a threelayered porous wall in all whorls,exceptforthelastone, whichhastwolayers. Stratigraphic data and age. Sample CG-31, Las Llacerias Formation, Castillo del Grajal section. Montiparus Zone, Kasimovian (lower Khamovnikian). Subfamily FUSULININAE von Möller, 1878 Genus Beedeina Galloway, 1933 Type species: Fusulina girty Dunbar & Condra, 1928 Beedeina cf. paradistenta (Safonova in Rauzer-Chernousova et al., 1951) Fig Measurements. L54.25 mm; D52.40 mm; L/D51.77; n57.5; D IV mm; d5 180 mm; wth540 mm. Remarks. For the rhombic short shell, number of volutions, and type of supplementary deposits, the studied specimen closely conforms to the characteristics of Beedeina paradistenta, from which it only differs in having a slightly larger L/D ratio. Stratigraphic data and age. Sample MZ-4, Picos de Europa Formation, Mazarrasa section. Upper part of the Fusulinella Zone, Fusulinella alvaradoi subzone, Moscovian (upper Myachkovian). Genus Pseudotriticites Putrja, 1940 Type species: Fusulina donbassica Putrja, 1939 Pseudotriticites? sp. 1 Fig Measurements. L57.50 mm; D52.50 mm; L/D53.0; n56.5; D IV mm; d5220 mm; wth555 mm. Remarks. Specimen showing a triangular shape, somewhat irregular septal folding and a three-layered wall pierced by pores. It mainly differs from the other Picos de Europa species assigned to Pseudotriticites by having a thicker wall and slightly curved axis of coiling. The somewhat irregular septal folding in some volutions makes its generic designation questionable, although the prominent central region suggests that it is closer to Pseudotriticites than to Quasifusulinoides. Stratigraphic data and age. Sample CG-16, Picos de Europa Formation, Castillo del Grajal section. Protriticites Zone, lower Kasimovian (Krevyakinian). Pseudotriticites cantabricus Villa sp. nov. Figs , 10.8, 10.11, 14.1 Beedeina? ex gr. acuta (Lee, 1927): van Ginkel, 1965, p , pl. 33, figs Pseudotriticites cf. lebedevi (Putrja, 1939): van Ginkel, 1971, p , pl. 5, figs Material. 7 axial sections, 1 nearly axial section, 5 slightly oblique sections, and numerous paraxial and oblique sections. Holotype: specimen CG-19/4a (Fig. 10.8). Derivatio nominis. From the names of the Cantabrian Mountains and the region of Cantabria, where the Castillo del Grajal peak section is located. Measurements. L mm; D mm; L/D ; n56 7; D IV mm; d mm; wth mm. Holotype: L57.95 mm; D52.55 mm; L/D53.12; n57; D IV mm; d5270 mm; wth5 50 mm. Description. Mature specimens with volutions exhibit subrhombic inner volutions and elongated fusiform outer shell, with blunt polar ends. Septa are regularly folded throughout the shell, usually showing rounded or subquadratic loops that do not reach the top of the chamber. Journal of Foraminiferal Research fora d 14/7/15 13:22:37 282

20 FUSULINES FROM THE PICOS DE EUROPA 283 FIGURE 14. Spirotheca of Pseudotriticites, Quasifusulinoides and Quasifusulina species from the Ándara sections. 1 Pseudotriticites cantabricus Villa sp. nov., CG-24/3 (in Fig. 10.3). 2 Pseudotriticites aff. oblonga (Nikitina, 1969) (in Fig ), PJ-0/5. 3 Quasifusulinoides parafusiformis Bensh, 1969, CG-27/4 (in Fig ). 4 Quasifusulinoides ex gr. parafusiformis Bensh, 1969, SCB-2/1 (in Fig. 11.7). 5 Quasifusulina sp. 1, AND-3/2b (in Fig ). 6 Quasifusulina praecursor Rauzer-Chernousova, 1951, AND-0/3 (in Fig ). Chomata are present in the inner volutions, but they are replaced by pseudochomata in the subsequent ones. Axial fillings are developed in early and intermediate volutions, varying from weak to rather strong, although never infilling the whole chamber. The wall, pierced by thin pores, is four-layered in the inner volutions and mostly three-layered (tectum, diaphanotheca, inner tectorium) in the outer ones (Fig. 14.1), although a thin outer tectorium may appear discontinuously in most specimens. Discussion. Although the specimens exhibit characters reminding us of several genera (Beedeina, Fusulina, Quasifusulinoides, Pseudotriticites), the shape of the inner volutions (usually rhombic, as in some Beedeina species) and pattern of septal folding (more regular than in Fusulina and Quasifusulinoides), along with the microstructure of the wall, suggest their belonging to Pseudotriticites Putrja, 1940, which, as pointed by van Ginkel (1971), seems to be a Beedeina-descendant genus. Pseudotriticites cantabricus sp. nov. recalls P. lebedevi Putrja, 1939, P. donbassicus (Putrja, 1939), and P. acutus (Lee, 1927). It differs from the three species in having larger size and stronger axial fillings; moreover, P. donbassicus shows stretched polar ends and P. acutus exhibits tighter coiling and smaller proloculus. Pseudotriticites asiaticus (Bensh, 1972), a species initially described from Fergana and also found in the Carnic Alps (Forke & Samankassou, 2000), recalls P. cantabricus sp. nov. in the overall morphology of the shell but it clearly differs by having much weaker axial fillings. Pseudotriticites cantabricus sp. nov. is probably conspecific with the material from the Corros Limestone (northern Palencia, southeast of Picos de Europa) initially described by van Ginkel (1965) as Beedeina? ex gr. acuta (Lee, 1927) and later by van Ginkel (1971) as Pseudotriticites cf. lebedevi (Putrja, 1939). Stratigraphic data and age. Samples CG-12, CG-14, CG- 16, CG-19, CG-24, Picos de Europa Formation, Castillo del Grajal section. Protriticites Zone, from the Moscovian/ Kasimovian transition interval to the lower Kasimovian (Krevyakinian). Pseudotriticites aff. oblonga (Nikitina, 1969) Figs. 10.7, , , 14.2 Measurements. L mm; D mm; L/D ; n ; D IV mm; d mm; wth mm. Remarks. Shape and size of the shell show high variability, the outer shape varying from rhombic (e.g., specimen PJ-0/11, Fig. 10.9) to elongated fusiform (specimen PJ-0/5, Fig ); inner whorls are usually fusiform, more rarely tending to rhombic. Septal folding intense and somewhat irregular; septal loops are usually rounded, sometimes subquadratic. Chomata present in several inner volutions, replaced by pseudochomata in the subsequent ones. Axial fillings weak and discontinuous. Wall four-layered, with gray diaphanotheca, and an inner tectorium much thicker than the outer one (Fig. 14.2); all layers are pierced by thin pores. This species resembles Pseudotriticites oblonga, which was originally described from the Primorsky Krai and assigned to Fusulina (Quasifusulinoides). The wall microstructure, septal folding, shape of shell, and type of Journal of Foraminiferal Research fora d 14/7/15 13:22:37 283

21 284 VILLA AND OTHERS chomata of the Picos de Europa species conforms well to the characteristics of the Primorsky Krai form. However, the scarcity of the Nikitina s (1969) material impedes evaluating the actual variability of P. oblonga. After obtaining such information, we could also estimate if Pseudotriticites turgida (Nikitina, 1969), described from the same locality, should be included in P. oblonga. Stratigraphic data and age. Samples MZ-10, Mazarrasa section, and PJ-0, Picos del Jierro section. Both from the Las Llacerias Formation. Protriticites Zone, Kasimovian (upper Krevyakian). Pseudotriticites sp. 2 Fig Measurements. L56.30 mm; D52.35 mm; L/D52.68; n56.5; D IV mm; d5260 mm; wth550 mm. Remarks. Shell showing subrhombic shape in section and intense septal folding. Chomata are present on the proloculus and the first three and a half volutions, being replaced by pseudochomata in the subsequent whorls. Axial fillings very weak, present only in a narrow belt along the axial region of the inner whorls. Wall pierced by pores and consisting of three layers: tectum, gray diaphanotheca and inner tectorium. This species somewhat resembles Pseudotriticites oblonga, but differs in its slightly shorter L/D ratio and the absence of an outer tectorium. Stratigraphic data and age. Sample AND-3, Las Llacerias Formation, Grajales Ridge locality. Montiparus Zone, Kasimovian (Khamovnikian). Genus Quasifusulinoides Rauzer-Chernousova & Rozovskaja in Miklukho-Maklay et al., 1959 Type species: Pseudotriticites fusiformis Rozovskaja, 1952 Quasifusulinoides? sp. 1 Figs , 11.6 Measurements. L mm; D mm; L/D ; n5 5 6; D IV mm; d mm; wth mm. Remarks. Main diagnostic features in this species are the elongated fusiform to subcylindrical test, pointed polar ends, and a usually large proloculus of irregular shape. Chomata are absent or restricted to two small tubercula on the proloculus. Septal folding is intense, usually forming archs or flat-topped loops. The wall is very thin and consists of tectum, gray diaphanotheca, and a discontinuous, thin inner tectorium; all these layers are pierced by weakly developed pores. Axial fillings can reach the top of the chambers in the axial region of some intermediate volutions. The generic assignment of this species to Quasifusulinoides is questioned in view of the fact that the thin wall, lack of chomata, and large and irregular proloculus are features more typical of Quasifusulina, a genus characterized by a two-layered wall. However, it also must be noted that primitive forms of the genus (e.g., Q. longissima praecursor Rauzer-Chernoussova in Rauzer-Chernousova et al., 1951) may exhibit a three-layered wall. All Quasifusulinoides species described in this paper also share an extremely thin wall, thus introducing some uncertainty in their generic assignment. However, this uncertainty is greater in Quasifusulinoides? sp. 1 due to the large and irregular proloculus of the present form. It resembles Q. longissima praecursor in the microstructure of the wall, L/D ratio, septal folding and extent of the axial fillings, but differs in having larger and more irregular proloculus and more acute polar ends. Stratigraphic data and age. Samples CG-14, CG-15, CG- 16, uppermost part of the Picos de Europa Formation, Castillo del Grajal section. Protriticites Zone, Kasimovian (lower Krevyakinian). Quasifusulinoides sp. 2 Fig Measurements. L56.55 mm; D51.80 mm; L/D53.64; n55.5; D IV mm; d5300 mm; wth540 mm. Remarks. This specimen occurs along with those assigned to Quasifusulinoides? sp. 1 in this study. Particularly, both forms share the similar microstructure of the wall. However, the regular rhombic shape of the inner whorls and the regular spherical proloculus of Q. sp. 2 seem to indicate that it is a different form from Q.? sp. 1. Stratigraphic data and age. Sample CG-15, uppermost part of the Picos de Europa Formation, Castillo del Grajal section. Protriticites Zone, Kasimovian (lower Krevyakinian). Quasifusulinoides parafusiformis Bensh, 1969 Figs , 14.3 Quasifusulinoides parafusiformis Bensh, 1969, p , pl. 12, fig. 6, pl. 13, figs Measurements. L mm; D mm; L/D ; n ; D IV mm; d mm; wth540 45mm. Remarks. Shell elongated fusiform throughout the growth, with rather flat median region and blunt-pointed polar ends. Wall essentially consisting of two layers plus an extremely thin and discontinuous lining that perhaps represents a vestigial inner tectorium (Fig. 14.3); these layers are pierced by weak pores. Chomata developed only on the proloculus. Septal folding moderately irregular and high. The specimens conform closely to those of Q. parafusiformis and, as in the types, they show a poorly-developed inner tectorium that suggests a possible link to Quasifusulina. Quasifusulinoides sp. 2, described above from the same section (sample CG-15) resembles present species, although it differs in exhibiting a more rhombic shape of the whorls. Stratigraphic data and age. Sample CG-27, top of the Picos de Europa Formation in the Castillo del Grajal section. Protriticites Zone, Kasimovian (Krevyakinian). Quasifusulinoides ex gr. parafusiformis Bensh, 1969 Figs , 11.13, 14.4 Remarks. Unfortunately, most of the specimens in this collection are broken and abraded, which hampers precise measurements. Nevertheless, it may be observed that although the shells are highly variable in length, from approximately 6 mm to more than 11 mm (Figs and 11.9), they exhibit rather similar values for the outer diameter ( mm) Journal of Foraminiferal Research fora d 14/7/15 13:22:38 284

22 FUSULINES FROM THE PICOS DE EUROPA 285 and number of whorls ( ), except for a microspheric specimen (Fig. 11.9) having at least nine volutions. Diameter of proloculus in macrospheric specimens is mm. Septal folding somewhat irregular and moderately high. Chomata developed only on the proloculus and the first volution. Axial fillings moderately developed. Wall very thin for the shell size, often consisting only of tectum and a gray diaphanotheca that is sometimes underlined by a thin and discontinuous inner tectorium (Fig. 14.4); the wall is pierced by weak pores. As in the form described above, these specimens also closely resemble those of typical Q. parafusiformis but differ in having, on average, a larger shell and smaller proloculus. The microspheric specimen exhibits a somewhat curved axis and reminds us of Quasifusulinoides sp. illustrated by Bensh (1969, pl. 13, fig. 5). Stratigraphic data and age. Sample SCB-2, SCB-3, top of the Picos de Europa Formation, Silla de Caballo Bajero section. Protriticites Zone, Kasimovian (Krevyakinian). Genus Quasifusulina Chen, 1934 Type species: Fusulina longissima von Möller, Quasifusulina praecursor Rauzer-Chernousova in Rauzer- Chernousova et al., 1951 Figs , 14.6 Quasifusulina longissima praecursor Rauzer-Chernousova in Rauzer-Chernousova et al., 1951, p. 325, pl. 58, figs Measurements. L mm; D52.50 mm; L/D54.22; n57; D IV mm; d5310 mm; wth555 mm. Remarks. The two Quasifusulina specimens found in this sample, of which one is illustrated and measured, fully match the characteristics of the forms described from the Sivin River area (Republic of Mordovia, Russia) by Rauzer-Chernousova in Rauzer-Chernousova et al. (1951) as a variety of Q. longissima (von Möller, 1878). The wall is relatively primitive for this genus due to a discontinuous and very thin inner tectorium (Fig. 14.6). This feature, as well as the presence of weak chomata on the proloculus and first volution, are the most distinctive characteristics of the species. Stratigraphic data and age. Sample AND-0, Las Llacerias Formation, collected from a locality at the Grajales Ridge situated close to Grajal de Arriba peak. Montiparus Zone, Kasimovian (Khamovnikian). Quasifusulina sp. 1 Figs , 14.5 Measurements. L57.50 mm; D52.00 mm; L/D53.75; n55; D IV mm; d5330 mm; wth535 mm. Remarks. This single specimen shows a slender shell coiled around a slightly bent axis of coiling. Septal folding clearly decreases in height towards the median region of the shell. Chomata are absent, except for two small and irregular tubercula on the proloculus. The wall is very thin, consisting of two layers pierced by distinct pores (Fig. 14.5). Although these characteristics do not conform to those of the known species, the scarcity of the available material prevents us from describing it as a new form. Stratigraphic data and age. Sample AND-3, Las Llacerias Formation, Grajales Ridge, in a locality close to La Rasa peak. Montiparus Zone, Kasimovian (Khamovnikian). Quasifusulina sp. 2 Fig Measurements. L57.70 mm; D52.50 mm; L/D53.08; n56; D IV mm; d5340 mm; wth550 mm. Remarks. Single specimen exhibiting a comparatively shorter shell compared to what is most common in this genus. Other species with smaller L/D ratio, somewhat resembling the present one, are Quasifusulina tenuis and Q. compacta, both described by Lee (1927). Quasifusulina tenuis is the most similar, differing only in having slightly stronger axial fillings, whereas Q. compacta, in addition to also having stronger axial fillings, shows a larger and more irregular proloculus. Also remarkable is the similarity of part of the original material from the Pamirs of Quasifusulina pseudotenuissima described by Leven & Davydov (2001, pl. 2, fig. 2) to Quasifusulina sp. 2. Stratigraphic data and age. Sample AND-3, Las Llacerias Formation, Grajales Ridge, in a locality close to La Rasa peak. Montiparus Zone, Kasimovian (Khamovnikian). Family SCHWAGERINIDAE Dunbar & Henbest, 1930 Genus Montiparus Rozovskaja, 1948 Type species: Alveolina montipara Ehrenberg, 1854, emend. von Möller, 1878 Montiparus ex gr. umbonoplicatus (Rauzer-Chernousova & Beljaev in Rauzer-Chernousova & Fursenko, 1937) Fig Measurements. L.6.10 mm; D52.50 mm; L/D.2.40; n56.5; D IV mm; d5130 mm; wth mm. Remarks. This single specimen deserves to be mentioned as it represents the first Montiparus occurrence in the Castillo del Grajal section. In spite of its poor orientation, the inferred shell shape and size, along with its wall microstructure (three layers in the inner volutions, including a thick outer tectorium; tectum, an incipient keriotheca, and a very thin, discontinuous inner tectorium in the outer ones) suggest a close relationship with Montiparus ex gr. umbonoplicatus reported by van Ginkel and Villa (1999) from the Las Llacerias section of the Cantabrian Mountains. This specimen could be conspecific with Montiparus aff. umbonoplicatus described below, but unfortunately its poor preservation hampers a proper comparison. Stratigraphic data and age. Sample CG-28, Las Llacerias Formation, Castillo del Grajal section. Montiparus Zone, Kasimovian (Khamovnikian). Montiparus aff. umbonoplicatus (Rauzer-Chernousova & Beljaev in Rauzer-Chernousova & Fursenko, 1937) Figs. 12.1, 12.3, 13.7 Measurements. L mm; D mm; L/D ; n56.5 7; D IV mm; d mm; wth mm. Remarks. The species conforms with the characteristics of M. umbonoplicatus in major diagnostic features such as the inflated central part of the shell, elongated and twisted polar ends, and moderate to weak septal folding. However, it differs from the latter in having, on average, a slightly larger size, smaller proloculus and somewhat broader chomata. Somewhat similar forms are M. subcrassulus Journal of Foraminiferal Research fora d 14/7/15 13:22:38 285

23 286 VILLA AND OTHERS (Rozovskaja, 1950) and M. sinuosus (Rozovskaja, 1950), which clearly differ from the Picos de Europa form in their stronger septal folding. Montiparus aff. umbonoplicatus differs from Montiparus cf. umbonoplicatus described below in having more volutions, larger L/D ratio on average, and a tighter spire (when comparing diameter of the 4th whorl in both species). Stratigraphic data and age. Sample AND-1, Las Llacerias Formation, Grajales Ridge localities. Montiparus Zone, Kasimovian (Khamovnikian). Montiparus cf. umbonoplicatus (Rauzer-Chernousova & Belajev in Rauzer-Chernousova & Fursenko, 1937) Figs. 12.4, Measurements. L mm; D mm; L/D ; n ; D IV mm; d mm; wth mm, isolatedly up to 90 mm. Remarks. In spite of the poor preservation of the outer shell of these specimens, which prevent a more precise identification, they can be considered as being very close, or identical, to Montiparus umbonoplicatus, with which they share similarity in size, type of chomata, septal folding, twisted polar ends (in some specimens), and a rather high variability in shell elongation. Stratigraphic data and age. Sample SCC-2, Las Llacerias Formation, Silla de Caballo Cimero section. Montiparus Zone, Kasimovian (Khamovnikian). Montiparus paramontiparus (Rozovskaja, 1950) Fig Triticites (Montiparus) paramontiparus Rozovskaja, 1950, p , pl. 1, figs Measurements. L mm; D mm; L/D ; n55 5.5; D IV mm; d mm; wth mm. Remarks. The two specimens available (one showing an ovoid shell and the other an inflated fusiform one, of which the latter is illustrated) conform to the variability in shell morphology observed in Triticites (Montiparus) paramontiparus paramontiparus Rozovskaja, The moderate size of the shell and the folded septa only in its polar ends also are in agreement with Rozovskaja s types. Stratigraphic data and age. Sample AND-1, Las Llacerias Formation, Grajales Ridge, in a locality close to Grajal de Arriba peak. Montiparus Zone, lower Kasimovian (Khamovnikian). Montiparus aff. fischeri van Ginkel, 1971 Figs. 12.6, , Measurements. L mm; D mm; L/D ; n ; D IV mm; d mm; wth mm. Remarks. Mature specimens show subcylindrical shell and massive chomata, which extend to the poles in the inner volutions and are broad and high, subquadrangular in section, in the subsequent ones. Septal fluting is present only in the polar ends. The wall, pierced by simple pores, is three-layered in the inner three-four volutions and two-layered in the last two whorls. Unpublished forms resembling Montiparus aff. fischeri have been found in other Picos de Europa localities, outside of the Ándara Massif, in a relatively short stratigraphic interval situated just below the Rauserites Zone. In this interval, specimens of the successive beds show a gradual but rapid development of the keriothecal wall and an increase of the septal folding, evidencing a probable transition to the genus Rauserites. Montiparus fischeri described from the Cabrales area (northern part of the Picos de Europa massifs) is considered as an advanced form in this Montiparus lineage because it shows a two-layered wall and a distinct keriotheca in the outer volutions. The present species from the Ándara Massif is very close to M. fischeri but differs in showing weaker septal folding and a more primitive wall, for the keriotheca is incipient and the outer tectorium is better developed (Figs ) than in the van Ginkel s (1971) species. Somewhat similar is Montiparus alaicus Bensh, 1972, from which Montiparus aff. fischeri can be distinguished by its weaker septal folding and much more massive chomata. Two specimens from Darvaz (Pamir) assigned to Montiparus sinuosus alaicus by Leven and Davydov (2001) also bear similarity to the Picos de Europa form. Stratigraphic data and age. Sample AND-2, Grajales Ridge, from a locality close to Grajal de Arriba peak, and samples SCC-2 and SCC-3, Silla de Caballo Cimero section, all samples from the Las Llacerias Formation. Montiparus Zone, lower Kasimovian (Khamovnikian). DISCUSSION CORRELATION OF THE ÁNDARA MASSIF STRATA WITH THE MOSCOW BASIN AND MOSCOVIAN/KASIMOVIAN BOUNDARY STRATOTYPE The Ándara Massif succession clearly contains the Moscovian/Kasimovian boundary, but fusuline provincialism during late Moscovian and early Kasimovian time hampers the precise correlation with the Moscow Basin boundary stratotype. During the Pennsylvanian these areas belonged to different fusuline biogeographic provinces. The Moscow Basin was situated in the Ural province (Rui et al., 1991) whereas the Cantabrian Zone was part of the western Paleo- Tethyan province (Villa et al., 2002). The neostratotypes for the Moscovian and the Kasimovian stages are recognized in the Moscow Basin succession (Makhlina et al., 2001a). The basal Kasimovian boundary horizon has been debated for years, but is currently placed at the base of the Suvorovo Formation in the Moscow Basin according to an official decision of the Interdepartmental Stratigraphic Committee (Rotai et al., 1978). An additional complication is that there appear to be several endemic fusuline morphotypes in the Cantabrian faunas, as shown in this study by the description of several new species and the frequent usage of the open nomenclature (i.e., aff. and ex gr.). These forms could not be confidently assigned to any known binomial name and might actually represent new species. Older beds sampled in the present research correspond to the upper (but not uppermost) part of the Picos de Europa Formation and are unequivocally of late Moscovian age. They contain abundant fusulines of the Fusulinella B Zone (van Ginkel, 1965), subzone of Fusulinella alvaradoi (5 subzone B3 of van Ginkel, 1965), which indicates Journal of Foraminiferal Research fora d 14/7/15 13:22:39 286

24 FUSULINES FROM THE PICOS DE EUROPA 287 FIGURE 15. Tentative correlation of the Ándara Massif succession with the Moscow and Donets basins units. a correlation with the middle member of the Peski Formation (upper part of the Myachkovian substage) of the Moscow Basin (Fig. 15). It must be noted, however, that the middle member of the Peski Formation is characterized by abundant species of the Fusulina cylindrica group, which are absent in the Ándara succession, and lacks species of the Fusulinella schwagerinoides group, which are typical of the upper Myachkovian in the Cantabrian Mountains. The composition of late Myachkovian fusuline assemblages of the Picos de Europa and other Cantabrian Zone areas show stronger affinities with Central Asian microfaunas, such as the Fusulinella species from the Alay Mountains described by Bogush (1963) and Dzhenchuraeva (1993). That correlation is established by the similarly advanced evolutionary stages of the Fusulinella species in both areas, and by taking into account the common presence of Fusulinella ex gr. pseudobocki. Overlying the Fusulinella alvaradoi subzone in the Picos de Europa succession, strata of the Protriticites Zone include the uppermost part of the Picos de Europa Formation and sequences 1 and 2 of the Las Llacerias Formation (Figs. 5 6). In the Moscow Basin, transitional forms between Fusulinella and Protriticites, assigned to primitive Protriticites by Russian authors, appear in the middle and upper members of the Peski Formation (Davydov, 1997, 2007; Isakova in Makhlina et al., 2001b; Isakova in Goreva et al., 2009). Those beds were included by Davydov (1997) in the Protriticites ovatus-praeobsoletes burkemensis zone of the uppermost Moscovian stage (Fig. 15). In the Ándara successions, Protricites ovatus Putrja, 1948 is absent, but specimens exhibiting wall characteristics similar to those of Praeobsoletes (here assigned to primitive Protriticites) occur in the lower part of the Protriticites Zone (samples CG-1 to CG-11 of the Castillo del Grajal section). Therefore, this part of the succession is tentatively considered to be of late Moscovian age. Above that, more typical Protriticites occur (e.g., Protriticites cf. manukalovae, P. sp. 2 ex gr. P. grosdilovae, P. grajalensis sp. nov.), but unfortunately none of them fully fit the characteristics of the few Protriticites specimens found in the Suvorovo Formation of the Moscow Basin (base of the Kasimovian stage). Therefore, in the Castillo del Grajal section, it is not possible to precisely place the Moscovian/Kasimovian boundary on the basis of the fusuline content, and instead only a tentative interval is indicated (Figs. 6 and 15). More advanced protriticitids occur near the top of the Picos de Europa Formation and in the lower part of the Las Llacerias Formation (sequences 1 and 2) including P. cf. subschwagerinoides, P. cf. mirabilis, and P. benshae sp. nov. (a species showing phrenothecae in which we include specimens from Central Asia described by Bensh, 1972); these occur along with large forms belonging to Quasifusulinoides and Pseudotriticites. Although the fusuline provincialism hampers a precise correlation with the Moscow Basin, the evolutionary stage of development of these forms indicates a probable equivalence with the interval from the Voskresensk Formation of the Moscow Basin (upper Krevyakinian), in which typical Protriticites occur (Heckel et al., 2007; Alekseev et al., 2009), up to the lower member of the Neverovo Formation (Khamovnikian) (Fig. 15). The basal member of the Neverovo Formation contains primitive Montiparus (Isakova in Alekseev et al., 2009), but this genus is not found below sequence 3 of the Las Llacerias Formation in the Picos de Europa succession. Therefore, the first appearance of Montiparus in the Picos de Europa succession is slightly younger than in the Moscow Basin (Fig. 15). It is possible that the gap in the basal part of the Las Llacerias sequence 3 is time-equivalent to the basal and lower Neverovo members or that sequence 1 of the Las Llacerias Formation is correlated with the mentioned Neverovo Formation members. The first appearance of Montiparus marked the base of the Montiparus Zone, which corresponds to sequence 3 of the Las Llacerias Formation. These beds are considered to be essentially equivalent to the middle and upper members Journal of Foraminiferal Research fora d 14/7/15 13:22:39 287