Selected Agglutinated Larger Foraminifera from the Font De Les Bagasses Unit (Lower Campanian, Southern Pyrenees)

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Carnets Geol. 15 (18)

Selected agglutinated larger foraminifera from the Font de les Bagasses unit (Lower Campanian, southern Pyrenees)

Sergi ALBRICH 1

Carme BOIX 2

Esmeralda CAUS 3

Abstract: The Font de les Bagasses marls and marly limestones (Montsec Mountains, NE Spain) host a rich assemblage of larger foraminifera, that was described for the first time in the early 20th Century from the currently submerged locality of Tragó de Noguera (Marginal Mountains). In this paper, a detailed architectural study has been carried out on selected, agglutinated larger foraminifera, which include Choffatella aff. rugoretis GENDROT, Hemicyclammina chalmasi (SCHLUMBERGER), Cuneolina conica ORBIGNY, Cuneolina cylindrica HENSON, and Dictyopsella kiliani MUNIER-CHALMAS. Moreover, it presents their biostratigraphic range and the range of the associated porcellaneous and lamellar-perforate for a- minifera in the study area. The assemblage is Early Campanian in age. Key Words: Larger foraminifera; systematics; biostratigraphy; Upper Cretaceous; Pyrenees; NE Spain. Citation: ALBRICH S., BOIX C. & CAUS E. (2015).- Selected agglutinated larger foraminifera from the Font de les Bagasses unit (Lower Campanian, southern Pyrenees).- Carnets Geol., Madrid, vol. 15, nº 18, p. 245-267. Résumé : Quelques grands foraminifères agglutinés de l'unité de la Font de les Bagasses (Campanien inférieur, Sud Pyrénées).- Les marnes et marno-calcaires de la Font de les Bagasses (Serra del Montsec, Nord-Est de l'Espagne) présentent une riche association de grands foraminifères décrite pour la première fois au début du 20e siècle dans la localité, aujourd'hui submergée, de Tragó de Noguera (Serres Marginals). Nous présentons ici une étude détaillée de l'architecture des grands foraminifères agglutinés : Choffatella aff. rugoretis GENDROT, Hemicyclammina chalmasi (SCHLUMBER- GER), Cuneolina conica ORBIGNY, Cuneolina cylindrica HENSON et Dictyopsella kiliani MUNIER-CHALMAS. Ce travail est complété par les répartitions stratigraphiques de ces espèces et par celles d'autres grands foramininifères, porcelanés et lamellaires-perforés, qui leur sont associés dans le secteur étudié. L'association est attribuée au Campanien inférieur. Mots-clefs : Grands foraminifères ; systématique ; biostratigraphie ; Crétacé supérieur ; Pyrénées ; NE Espagne.

1. Introduction for the study of the "Upper Santonian" larger foraminifera until the 1960s, when the locality The larger foraminifera of the southern Pyre- was submerged under the water of the Santa nees and particularly those attributed to the Late Anna reservoir. However, "Tragó de Noguera Cretaceous have been studied since the end of fauna" is well represented in the Font de les Ba- the 19th Century. The pioneer was the Spanish gasses Unit in the nearby Montsec Mountains engineer Luis Mariano VIDAL who, during the and since that time studies have continued there works for the first geological map of Lleida pro- (HOTTINGER 1966; HINTE, 1966; WANNIER, 1983; vince (NE of Spain), collected abundant material BOIX et al., 2009; HOTTINGER & CAUS, 2009; containing larger foraminifera. This material was ALBRICH et al., 2014, among others). However, a sent to the French naturalist M.C. SCHLUMBERGER detailed inventory on the larger foraminifer fau- who, in 1899, described various genera and spe- na has not been carried out yet. Some groups cies (e.g., Dictyopsella chalmasi, Dictyopsella such as lacazinids (HOTTINGER et al., 1989), kiliani, Vidalina hispanica, Nummofallotia creta- meandropsinids (HOTTINGER & CAUS, 2009) or cea, and Lacazina elongata) from the locality of orbitoidids (ALBRICH et al., 2014) have received Tragó de Noguera (Marginal Mountains, Lleida, the attention of researchers, while others lack Fig. 1). After SCHLUMBERGER's work, the so-called detailed studies and an accurate biostratigraphic "Tragó de Noguera fauna" became a reference distribution.

1 Departament de Geologia (Paleontologia), Universitat Autònoma de Barcelona, E-08193 Cerdanyola del Vallès (Bellaterra) (Spain) [email protected] 2 Repsol Campus, Méndez Álvaro 44, 28045 Madrid (Spain) 3 Departament de Geologia (Paleontologia), Universitat Autònoma de Barcelona, E-08193 Cerdanyola del Vallès (Bellaterra) (Spain) [email protected] Published online in final form (pdf) on November 18, 2015 [Editor: Bruno GRANIER; technical editor: Christian C. EMIG; language editor: John A. STARMER]

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Figure 1: Location of the study area (about 75 km N of Lleida, Catalonia). Note the position of the Santa Anna reservoir with the submerged old locality of Tragó de Noguera. The capital letters refer to the stratigraphic sections studied in this paper: A. Font de la Plata section; B. Terradets section; C. Clot d'Olsí section, D. Montrebei section.

Therefore, the aim of this paper is to contri- along the southern slope of the mountain, dip- bute to the knowledge of foraminifera from the ping 35º-40º to the North. They rest uncon- Font de les Bagasses Unit by presenting a detai- formably on the deposits of the La Cova Unit led architectural analysis of the main aggluti- (uppermost Lower Coniacian to Middle Santonian nated larger foraminifera and their stratigraphic according to BOIX et al., 2011), passing distribution. A tentative range chart including gradually upwards to the deposits of the agglutinated forms, together with the main lar- Terradets Unit (PONS, 1977; SIMÓ, 1986), with a ger porcellaneous and lamellar-perforate forami- base dated latest Early to Middle Campanian nifera, is also given as an useful tool for future (VILLALONGA, 2009). geological studies. Palaeogeographically, the Upper Cretaceous deposits of the Montsec Mountains were depo- 2. Geographical and Geological setting sited in the southern margin of the Pyrenean The Montsec Mountains extend continuously Basin that extended approximately E-W and E-W for more than 50 kilometers in the opened to the Atlantic Ocean (Bay of Biscay). Southern Pyrenees, between the Tremp and The basin was locaced at about 30ºN latitude Àger-Vilanova de Meià basins (Lleida, NE of (DERCOURT et al., 1985; HAY et al., 1999). The Spain). The Boix, Noguera Pallaresa and Nogue- climate during the Santonian-Campanian was ra Ribagorçana rivers, which run N-S, individua- warm (PUCÉAT et al., 2003) and the global sea- lize three adjacent geographic units, from East level curve indicates a level 50 m higher than to West: Montsec de Rúbies, Montsec d'Ares, the present day for the Early Campanian (MILLER and Montsec d'Estall (Fig. 1). et al., 2005). From a geological point of view, the Montsec The age "Late Santonian" traditionally attri- Mountains belong to the Montsec Thrust Sheet, a buted to the Font de les Bagasses deposits (HOT- Southern Pyrenean upper thrust sheet detached TINGER, 1966; HINTE, 1966; PONS, 1977; CAUS & from its basement over Upper Triassic evapo- CORNELLA, 1983; CAUS & GÓMEZ-GARRIDO, 1989a, rites. At the South, it shows a simple anticline 1989b; CAUS et al., 1999; among others) has structure representing the front of the Montsec been recently contested (ALBRICH et al., 2014). thrust, while northwards it develops a broad New data published in this last paper, obtained by syncline supporting a piggyback basin filled with means of isotope strontium stratigraphy, indica- Palaeogene rocks, the Tremp-Graus Basin te a younger age for the whole unit. Thus, the age (MUÑOZ, 1992; TEIXELL & MUÑOZ, 2000). The attributed to the larger foraminifera from the Font Montsec Mountains include a complete Mesozoic de les Bagasses Unit is Early Campanian, corres- sequence, mainly Upper Cretaceous, of about ponding to the ammonite zones of Placenticeras 1500 m in thickness (Fig. 2). tridorsatum and Menabites delawarensis, and to the planktonic foraminiferal zone of Globotrun- The deposits attributed to the Font de les canita elevata (see the correspondence in the Bagasses Unit ("Font de les Bagasses marls and global time scale of GRADSTEIN et al., 2012). clays" in PONS, 1977) uniformly extend E-W

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Figure 2: Simplified geological cross-section of the Montsec Thrust Sheet along the Montrebei pass (Noguera Ribagorçana River). Modified from TEIXELL & MUÑOZ (2000).

3. Methodology 307891.0 and Y: 4660320.0. Samples from this section are named ME-23 to The work is based on field observations and ME-40. 137 samples collected from four stratigraphic sections on the well-exposed slopes of the Ten samples are from washed sediments Montsec de Rúbies and Montsec d'Ares. From while 127 are from consolidated rocks. The East to West, the sections are (Figs. 3 - 4 - 5 - samples from unconsolidated sediments were 6): washed over two sieves with holes of 0.35 and A) Font de la Plata section measured in the 0.21 mm respectively, and larger foraminifera Font de la Plata gully about 2 km east of were picked, photographed and sectioned. The the Rúbies abandoned village; external features of the foraminifera shells were coordinates: UTM ETRS89 31T X: photographed using optical and scanning 328852.3 and Y: 4654008.6. Samples electron microscopes. Oriented sections were from this section are labelled SG-7 to prepared following the classical techniques for SG-43. studying internal characteristics in agglutinated B) Terradets section measured along the foraminifera (see HOTTINGER, 1967). Larger Noguera-Pallaresa River in the well- foraminifera pictures are at standard exposed trench of the C-13 road (a field magnifications of x25 and x50. More than 350 picture of this section is shown in thin-sections were obtained from hard samples ALBRICH et al., 2014, Fig. 4A); to study the larger foraminifera structures in coordinates: UTM ETRS89 31T X: random sections and the microfacies analysis. 324903.0 and Y: 4655864.0. Samples All the larger foraminifera presented in this from this section are named MARG-122 paper were identified at genus and species to MARG-177. levels. The genera were defined by the presence C) Clot d'Olsí section measured along the or absence of structural elements combined with Pardina gully near the Ager-Alçamora chamber arrangement, while the species are road; coordinates: UTM ETRS89 31T X: defined by quantitative morphological 311246.8 and Y: 4657201.0. Samples characteristics. from this section are labelled CO-28 to All the material studied in this work was CO-68. deposited in the collection of Paleontology of the D) Montrebei section measured along the Department of Geology (Universitat Autònoma left margin of the Noguera-Ribagorçana de Barcelona). The figured specimens are River in the Montrebei pass; labeled as PUAB 82410, 82412, and 84458- coordinates: UTM ETRS89 31T X: 84473.

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Figure 3: La Font de la Plata stratigraphic section with the larger foraminifera distribution.

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Figure 4: Terradets stratigraphic section with the larger foraminifera distribution.

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Figure 5: Clot d'Olsí stratigraphic section with the larger foraminifera distribution.

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Figure 6: Montrebei stratigraphic section with the larger foraminifera distribution.

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4. Stratigraphy and microfacies Based on field work, palaeonto- logical and sedimentological studies, the Font de les Bagasses Unit is divided into five stratigraphic subunits that are, from bottom to top (Figs. 3 - 4 - 5 - 6): 4. a. Subunit 1 This basal subunit, which overlies La Cova Unit by means of an erosive contact (CAUS et al., 1999), is composed of yellow sands, clays and coarsely grained sandstones with some intercalated micro- conglomerate beds; it is interpreted to be deposited in a fluvial setting. This subunit is well represented in the Montsec de Rúbies, with a maximum thickness of 20m in the Font de la Plata section, but disap- pears to the West of the Terradets section (Figs. 3 - 4). In the uppermost part of the subunit, some san- dy limestones with fragmented marine bivalves indicate the transition Figure 7: Microfacies of the subunit 2. A. Pelletoidal packstone with from fluvial to littoral marine porcellaneous larger foraminifera as major components; from SG-8. B. deposits (CLAPÉS, 1997). This sub- Peletoidal packstone-wackestone with agglutinated and porcellaneous unit lacks larger foraminifera. foraminifera; from SG-10. 4.b. Subunit 2 The subunit 2 (called "Boix River platform" in CAUS et al., 1999) overlies the previous subunit in the sections of Font de la Plata and Terradets, disappearing to the West. It is composed of well- bedded nodular limestones with pelletoidal-packstone texture containing a diverse fauna of gastropods, bivalves (including rudists), echinoids, larger foraminifera and some sponge spicules (Fig. 7 A-B). Dasycladacean algae occurring in this subunit are frequently recrystallized. The unit reaches a maximum thickness of 10 m in the Font de la Plata section and disap- pears west of the Terradets section (Pla d'Escumó area). Among the foraminifera, the porcellaneous Pseudolacazina cantabrica HOTTIN- GER et al. and Alexina papyracea HOTTINGER & CAUS are the main components of the facies associated with the agglutinated foraminifera (Figs. 3 - 4). Figure 8: Microfacies of the subunit 3. A. Wackestone-packstone with P. sphaeroidea, porcellaneous foraminifera, spicule sponges and fragmented dasycladacean algae (close to the scale bar: an oblique transverse section of Trinocladus tripolitanus RAINERI); from samples M-125. B. Wackestone with agglutinated foraminifera, small benthic and dasycladacean algae; from M-145.

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4.c. Subunit 3 In the Font de Plata and Terra- dets sections, the subunit 3 overlies the subunit 2 and in the Clot d'Olsí and Montrebei sections the subunit 3 overlies directly the deposits of the La Cova Unit (Figs. 3 - 4 - 5 - 6). It is composed of gray and yellow argillaceous marls intercalated with nodular marly limestone with pelletoidal-packstone texture, and more rarely wackestones (Fig. 8 A-B). Fine quartz grains are rare. In the Clot d'Olsí and Montrebei sections the subunit shows intra- formational slumps. The thickness varies from 135 to 210m. The faunal content is abundant and diverse: foraminifera, gastropods, bivalves (oysters, rudists and indeterminate bivalves), corals, bra- chiopods, and echinoid fragments are the main components (NOGUÉS, 2000). Gastropods, rudists and other indeterminate bivalves are frequently recrystallized. Sporadi- cally, green algae are also present in some levels, with fragments of Figure 9: Microfacies of the subunit 4. A. Sandstone with isometric crustose rhodophyte algae occur- irregular quartz grains, elongate bivalve fragments, some rotaliids and agglutinated foraminifera; from SG-40. B. Packstone with irregular quartz ring in the uppermost part of the grains, fragmented bryozoans, echinoids, bivalves, worms and lamellar- subunit. Larger foraminifera are perforate foraminifera; from Me-29. abundant and highly diverse (see their distribution in Figs. 3 - 4 - 5 - 6). 4.d. Subunit 4 The boundary between this subunit and the underlying subunit 3 is marked by the presence of coarse- grained quartz in the Font de les Bagasses deposits. However, the coarse terrigenous deposits are not uniformly distributed in the unit: in the Font de la Plata and Montrebei sections, discontinuous lenticular- shaped calcareous sandstones are interbedded within argillaceous marls and marly limestones in intervals of 50-to-100m (Figs. 3 - 4 - 5 - 6), while in the Terradets and Clot d'Olsí sections the terrigenous intervals do not exceed a few meters (Figs. 4 - 5). The fauna of invertebrates and larger foraminifera appear frequently fragmented (Fig. 9 A-B).

Figure 10: Microfacies of the subunit 5. A. peletoidal wackestone with sponge spicules, fragmented bryozoans and lamellar-perforate foraminifera; from M-170. B. Packstone-grainstone with crustose rhodophyte algae, bryozoans and echinoid fragments, and small foraminifera; from M- 167.

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4.e. Subunit 5 MINSKI's (2014) classification is only partially followed, and some remarks or new proposals This uppermost subunit is composed of grey are added to this classification (see below). or yellow marls and marly limestones alternating Finally, the hierarchy of taxonomic criteria sket- with nodular and, more rarely, bioclastic lime- ched by HOTTINGER (1978, Table 1) has been stones. The microfacies study shows wacke- taken into account in the elaboration of our pro- stone, packstone and grainstone textures (Fig. posals. In order to facilitate the comparison bet- 10 A-B). Rarely, coarse-grained idiomorphic ween the architectural characteristics of the ge- quartz grains are present. Bryozoans and spon- nera of agglutinated foraminifera described in ge spicules are the dominant components of the this paper, the architectural and structural terms marly limestones associated mainly with large are those referred to by HOTTINGER (2006). specimens of Larrazetia larrazetti MUNIER-CHAL- MAS. The nodular limestones contain brachio- Phylum Foraminifera pods, rudists and other bivalves, and echino- (ORBIGNY, 1826) derms; the larger foraminiferal fauna is domi- PAWLOWSKI et al., 2013 nated by Lacazina elongata MUNIER-CHALMAS. In constrast, the bioclastic limestones include Class Globothalamea mainly Praestorsella roestae (VISSER), Praesi- PAWLOWSKI et al., 2013 derolites praevidali (NEUMANN), and Pararotalia Order Loftusiida tuberculifera (REUSS). The limestones of the KAMINSKI & MIKHALEVICH Terradets Unit conformably overlie this subunit. in KAMINSKI, 2004 5. Systematic micropaleontology Remarks: Following the authors of the Order In the taxonomic classification of larger Loftusiida, it includes all agglutinated forami- foraminifera, it is commonly accepted that nifera with coiled arrangement of chambers in genera are identified by the presence or absence early stage, that later may be uncoiled, and with of shell structures like exoskeleton, endo- complex exoskeleton. The simple forms with the skeleton and supplemental skeleton combined same chamber arrangement are grouped in the with the chamber arrangement. Species are Order Lituolida LANKESTER, 1885. Therefore, in related mainly to the adult size and the size of the future, further studies are needed to explore chamber elements (HOTTINGER, 1978; CAUS, the relationship between taxa included in both 1981), although a great number of oriented and orders. random sections have been required for Suborder Loftusiina correctly interpreting the shell structures and KAMINSKI & MIKHALEVICH comparing the biometric data of the shells at in KAMINSKI, 2004 homologous growth stages. The ascription of genera to taxonomically higher-levels is contro- Remarks: KAMINSKI (2014) refers to the versial due to the lack of a comprehensive Suborder Loftusiina as "test free or attached, systematic classification based on phylogenetic multilocular, coiling or uncoiling; wall aggluti- characteristics. As a result, the successive clas- nated, microgranular, with imperforate epider- sifications of LOEBLICH & TAPPAN (1964; 1988; mis (outer layer) and an inner alveolar layer; 1992; 1994) have been mainly used in for a- aperture terminal in the initial stage, later mul- miniferal works, frequently without discussion. tiple, rows of rounded areal openings". There- In recent years, the integration of molecular fore, with reference to the order description, the data in foraminiferal studies has allowed the type and distribution of the apertures are added. highest-levels foraminiferal classification (see for Observations: In KAMINSKI (2014) the term instance, PAWLOWSKI et al., 2013, and the papers "epidermis" is used in a wider meaning than in cited therein). For categories below the class, its original description (HENSON, 1948), which multiple classifications are proposed, but only a restricted the term epidermis for walls with poly- few of these are clearly presented, as the hie- gonal subepidermal network (HOTTINGER, 1967). rarchy of taxonomic criteria used for such classi- The inner alveolar layer of KAMINSKI (2014) fications remains obscure and is often incon- refers to the "alveolar network" of DOUVILLÉ sistent. For the agglutinated foraminifera, KA- (1906) comprising the alveolar and subepider- MINSKI (2014) updated a new classification par- mal exoskeleton types of HOTTINGER (1967). The tially based on the LOEBICH and TAPPAN (1988; former consists of a tapestry of one or several 1992; 1994) and KAMINSKI (2004) classifications. generations of rounded outline alveoli, with no However, some criteria proposed by KAMINSKI differentiation between beams and rafters. The (2014) for grouping larger, complex foraminifera latter has differentiated beams and rafters for- are only partially defined. Thus, in this paper the ming a regular polygonal subepidermal network phylum and class categories proposed by similar to the pigeonholes. The thin epidermis is PAWLOWSKI et al. (2013) are accepted, but KA- restricted to this last group of exoskeleton type.

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Superfamily Loftusoidea BRADY, 1884 ments and the communication system of apertures. Remarks: Foraminifera with planispiral (or streptospiral) arrangement of chambers that Subfamily Choffatellinae MAYNC, 1958 may become uncoiled, peneropliform or cyclical Remarks: In this work, the Subfamily Chof- in the adult stage, with exoskeleton structures fatellinae, included by KAMINSKI (2014) in the and one or several apertures. Family Cyclamminidae, is removed from this last KAMINSKI (2014) included the following four family and transferred back to Spirocyclinidae. families in this superfamily: the Jurassic Meso- Accordingly to MAYNC (1958), it is characterized endothyridae with a pseudokeriothecal exoske- by planispiral to uncoiled or peneropliform-to- leton (see SEPTFONTAINE, 1988, for details); the cyclical shells with polygonal subepidermal exo- Late Jurassic to recent Cyclamminidae with an skeleton, multiple aligned apertures, and lacking alveolar exoskeleton (sensu HOTTINGER, 1967); endoskeletal elements. and the Jurassic-Cretaceous Spirocyclinidae with Choffatella SCHLUMBERGER, 1905 polygonal subepidermal exoskeleton. The fourth Family Loftusiidae has the same structural cha- (Type-species: C. decipiens SCHLUMBERGER, racteristics as Spirocyclinidae, but the axis of 1905) coiling is elongated. The Family Haurinidae SEPT- Description: Agglutinated, lens-compressed FONTAINE, 1988, with similar disposition of cham- shell with multiple openings placed in one row in bers and exoskeleton structures, has been ascri- the median plane of the apertural face. The bed to the Suborder Orbitolina by KAMINSKI chambers are planispiral-evolute arranged. (2004, 2014) (see the discussion below). Choffatella has exoskeleton elements (beams Family Spirocyclinidae and rafters) forming a well-developed polygonal subepidermal network. It lacks endoskeleton, (MUNIER-CHALMAS, 1887) MAYNC, 1950 although LOEBLICH & TAPPAN (1988) talked about Remarks: The Family Spirocyclinidae was an "endoskeleton formed by thick massive sep- constructed by MUNIER-CHALMAS (1887) to include ta". The septa are pierced by multiple foramina four agglutinated genera of larger foraminifera aligned from one septum to the next. (Spirocyclina, Dicyclina, Cuneolina and Orbito- Age: After LOEBLICH & TAPPAN (1988), the lina), having in common exoskeleton elements genus Choffatella ranges from the Late Jurassic forming a polygonal subepidermal network, but to the Late Cretaceous (Cenomanian). GENDROT with a different chamber arrangement. (1968) and SEPTFONTAINE (1980) enlarged its MAYNC (1950) emended the Spirocyclinidae, stratigraphic range to the Santonian. Our speci- resulting in the following description: planispiral- mens attributed to this genus are collected in to-peneropliform or uncoiled agglutinated shells deposits attributed to the Lower Campanian with one or multiple apertures. Therefore, the (ALBRICH et al., 2014). biserial genera Dicyclina and Cuneolina and the Choffatella aff. rugoretis GENDROT, 1968 uniserial Orbitolina were removed from Spiro- cyclinidae. (Fig. 11 A-F) Synonymies: All representatives of the Spirocyclinidae have an exoskeleton constituted by beams and aff. 1968 Choffatella rugoretis GENDROT, Pl. 4, figs. 6- rafters forming a polygonal subepidermal net- 13. aff. 1972 Gendrotella rugoretis (GENDROT) MAYNC, Figs. work covered by a fine epidermis. According to 5-6. AYNC M (1950), both taxa, with or without exo- 1977 Choffatella rugoretis (GENDROT) CORNELLA, Fig. 3. skeleton elements, were included in Spirocy- 1978 Choffatella rugoretis GENDROT, CAUS et al., Fig. 2. clinidae. But LOEBLICH & TAPPAN (1988), not in Description: Finely agglutinated, lens- accordance with MAYNC (1950), restricted the Fa- shaped foraminifera with one row of openings mily Spirocyclinidae to the genera with endo- aligned in the median plane of the apertural skeleton elements, while most of the genera face. The A-generation consists of an embryo lacking such elements were placed in the Family formed by a spherical protoconch and a hemi- Cyclamminidae. The same classification was also spherical deuteroconch followed by planispiral- followed by KAMINSKI (2014). However, Cyclam- evolute chambers disposed in two or two and mina (type-genus of the Family Cyclamminidae) half whorls. The earliest chambers have a qua- possesses alveolar exoskeleton and not a regular drangular outline, but they become rapidly low polygonal subepidermal network as in Spirocycli- and elongate to build an operculiniform spire. nidae. Therefore, the two families should remain The number of chambers in the last whorl varies separate, and the genera with polygonal subepi- from 24 to 27. The septa are arched and pierced dermal patterns should be ascribed to the Family by multiple, regularly distributed intercameral Spirocyclinidae instead of the Cyclamminidae. foramina. The average size of the adult mega- Following HOTTINGER (1978), the presence or ab- lospheric shells is 2.6 mm. The scarce and sence of endoskeleton elements is considered poorly preserved B-forms can reach 3.0 mm and here of lower category than the exoskeleton ele- up to three whorls of planispiral chambers.

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Figure 11: A-C. Choffatella aff. rugoretis: subequatorial sections (near the equatorial plane) showing the planispiral, partially evolute chamber arrangement and multiple aligned intercameral foramina; from CO-32. D-F. Choffatella aff. rugoretis: lateral view showing the elongated chambers and sutures (scanning microphotography); the epidermis is poorly preserved. from SG-36. G-I: Hemicyclammina chalmasi:G-H; Hemicyclammina chalmasi: external views showing the inflate chambers and depressed sutures (scanning microphotography); from SG-36. I. Hemicyclammina chalmasi: equatorial section of an extraordinarily well preserved specimen showing the large intercameral foramen and its particular position in the septum (hemi-septum from MAYNC, 1953); from SG-36. Scale bar 1 mm.

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Remarks: MAYNC (1972) re-studied the GEN- 1953). Hemicyclammina has a well-developed DROT's types of C. rugoretis and he concludes polygonal subepidermal network with two orders that the species attributed by GENDROT (1968) to of beams. It lacks endoskeleton. the genus Choffatella lacked an exoskeleton. Age: After LOEBLICH & TAPPAN (1988), the age Thus, MAYNC (1972) stated that GENDROT's taxon of Hemicyclammina is Early Cretaceous and cannot be assigned to the genus Choffatella, a Cenomanian for this genus but, with the Jurassic-Early Cretaceous genus exhibiting an ascription of "Dictyopsella" chalmasi to this genus exoskeleton consisting of beams and rafters or- (CAUS et al., 1979), the range extends as far as ganized in a polygonal subepidermal network. the Early Campanian. Therefore, he erected the new genus Gendrotella for it. Hemicyclammina chalmasi (SCHLUMBERGER, 1899) However, material from the type locality (Martigues, Bouches-du-Rhône, South-East of (Fig. 11 G-I) France) has yielded specimens morphologically Synonymies: identical to those described by GENDROT (1968) 1899 Dictyopsella chalmasi SCHLUMBERGER, Pl. 8, fig. 4. and bearing exoskeleton elements; moreover, 1977 Hemicyclammina chalmasi (SCHLUMBERGER) the specimens of C. rugoretis showed by GEN- CORNELLA, Figs. 4-5. DROT (1968, Pl. 4, figs. 9-10) exhibit short 1978 Hemicyclammina chalmasi (SCHLUMBERGER) CAUS et beams and rafters forming a delicate network in al., Figs. 1- 5. the inner surface of the chamber, interpreted cf. 1999 Hemicyclammina chalmasi (SCHLUMBERGER) here as a subepidermal type of exoskeleton. POLAVDER, Pl. 1, figs. 1-6. cf. 2004 Hemicyclammina chalmasi (SCHLUMBERGER) Thus, the species is re-installed within the genus SCHLAGINTWEIT & WAGREICH Pl.1, figs. 1-9. Choffatella, and Gendrotella becomes a non- cf. 2010 Hemicyclammina chalmasi (SCHLUMBERGER) valid genus after the ICNZ. RADOIČIĆ et al., Pl. 1, figs. 1-5. Similarities and differences: The speci- Description: Coarsely agglutinated, lens- mens from the Font de les Bagasses Unit are shaped shells with a more or less rounded peri- larger (almost the double size) than the size of phery. The diameter varies from 1.3 to 1.5 mm. Martigues (type-locality of C. rugoretis), and de- The slightly inflated chambers have a planispiral- signated as Choffatella aff. rugoretis. Further involute arrangment, with two-to-three spiral studies are needed to determine if the size diffe- whorls. The sutures are depressed, exhibiting a rence is due to evolution (Santonian in the GEN- lobate outline. Two generations have been DROT's specimens and Early Campanian in the observed, interpreted as A and B generations Pyrenean specimens) or related to a deeper (CAUS et al., 1978, Figs. 1-4), although the adult water environment in the Font de les Bagasses size does not present any differences. The A- Unit than in the Martigues deposits. generation consists of a spherical embryo Stratigraphical distribution: In the Font de followed by two whorls of planispiral chambers, les Bagasses, C. aff. rugoretis occurs in the four which generally number 8 or 9 in the last whorl, measured sections. It has been identified mainly but some specimens can reach 11 chambers. In in the washed sediments, but some random the B-generation, the small microsphere is follo- sections also occur in the consolidated rocks. wed by three whorls of spiral-involute chambers See the distribution in Figs. 3 - 4 - 5 - 6. with 6-8 chambers in the last whorl. The septa are oblique. The exoskeleton is well developed Geographical distribution: It has been with long beams and less pronounced rafters, but only mentioned from the Pyrenean basin (North can only be observed in the thin-sections of the of Spain and South and South-East of France). isolated specimens (Fig. 13 I). The Hemi- Subfamily Hemicyclammininae cyclammina sections coming from hard rocks BANNER, 1966 (Fig. 13 A-F) are difficult to identify and are designated in this work as H. cf. chalmasi. Remarks: Agglutinated foraminifera with po- However, they coincide with the sections shown lygonal subepidermal exoskeleton. The cham- by POLAVDER (1999), SCHLAGINWEIT & WAGREICH bers with planispiral-involute arrangement have (2004) and RADOIČIĆ et al. (2010). only one aperture. It lacks endoskeletal ele- Similarities and differences: The specimens ments. of the Font de les Bagasses Unit are identical to Hemicyclammina MAYNC, 1953 those described by SCHLUMBERGER (1899) from the Tragó de Noguera under the name of Dictyopsella (Type-species: Hemicyclammina sigali MAYNC, 1953) chalmasi, that have been transferred to the genus Hemicyclammina by CAUS et al., 1978. Description: Agglutinated, lens-shaped fora- Hemicyclammina chalmasi differs from H. minifer with rounded-to-subacute periphery. The sigali MAYNC (species-type of the genus) due to its chambers have a planispiral-involute arrang- larger size, the complexity of the skeleton, and ment. The aperture is unique, large, located the septa perpendicular to the external wall. near the base of the septum and with circular- to-elliptic outline (hemi-septum after MAYNC,

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Stratigraphical distribution: The age of H. Remarks: Although the "plexus" Pseudo- chalmasi is Early Campanian. In the Font de les cyclammina (SEPTFONTAINE, 1980; 1988) is men- Bagasses Unit, it is observed in all sections, al- tioned as having a long stratigraphic extension, though infrequently. The sections from consoli- the idea of a polyphyletic group is recurring in the dated rocks are badly preserved. literature (see SEPTFONTAINE, 1980, and the papers Geographical distribution: It has been cited therein). BANNER (1970) placed the genus mentioned in the Pyrenees and the central Euro- Pseudocyclammina together with Choffatella in pe (SCHLAGINWEIT & WAGREICH, 2004) and Balkans the Choffatellinae; however, both genera differ in (POLAVDER, 1999; RADOIČIĆ et al., 2010). the foraminal pattern: aligned in Choffatella and cribrate in Pseudocyclammina. Subfamily Amijiellinae Pseudocyclammina sphaeroidea SEPTFONTAINE, 1988 GENDROT, 1968 Remarks: After SEPTFONTAINE (1988), the (Fig. 12 A-L) Subfamily Amijiellinae consists of planispiral to uncoiled shells with polygonal subepidermal exo- Synonymies: skeleton, multiple cribrate apertures, and 1968 Pseudocyclammina sphaeroidea, GENDROT, Figs. 1- lacking endoskeletal elements. SEPTFONTAINE 5. (1988) and also KAMINSKI (2014) included the 2013 Pseudocyclammina sphaeroidea, CAUS et al., Fig. Subfamily Amijiellinae in the Family Haurinidae 7g. Description: Small and globular Pseudocy- (SEPTFONTAINE, 1988), which is characterized by clammina (the observed diameter varies from 0.6 planispiral to peneropliform shells with a poly- mm to 1.1 mm). Dimorphism is not observed in gonal subepidermal (hypodermal in SEPTFON- our material from the Font de les Bagasses: all TAINE, 1980) network. However, such characte- the specimens have similar size at the same ristics are shared with the Family Spirocyclinidae growth stage. Where visible, a small and (see above), which has priority. However, fur- spherical protoconch is followed by a small hemi- ther studies are needed to clarify the relation- spherical deuteroconch. The post-embryonic ship between the two families: Haurinidae used chambers exhibit a planispiral-involute arrang- mainly for the Jurassic taxa (see the excellent ment, resulting in two and half to three whorls of work of SEPTFONTAINE, 1980, 1988), and Spirocy- spire, rarely three and half whorls. Specimens clinidae for the Cretaceous taxa. showing uncoiled stage are not frequent in our In addition, KAMINSKI (2014) placed the Sub- material. The earliest chambers are quadrangular family Amijiellinae (and the Family Haurinidae) in shape, but in the latest stage the chambers in the Suborder Orbitolinina KAMINSKI, 2004, that become low and elongated with peneropliform was defined as "trochospiral to conical, later morphology. The number of chambers in the last stage may have reduced number of chambers whorl varies from 10-12 in the small forms to 13- per whorl, or may become uniserial and recti- 14 in the largest ones. The external wall linear". However, neither the Subfamily Ami- (epidermis) of our specimens is diagenetically jiellinae nor the Family Haurinidae possesses altered. Moreover, the specimens incorporate such characteristics and, in the current state of angular, coarse quartz grains in the external wall knowledge, they should be removed from Orbi- and in the septa, frequently masking the tolinina and included in Loftusiina. They are exoskeleton structures. Where they are visible; presented in this paper this way. these structures show short beams and rafters Pseudocyclammina forming a poorly developed subepidermal YABE & HANZAWA, 1926 network. The chamber lumen is free of endoskeletal elements. The septa are massive and (Type-species: Cyclammina lituus YOKOHAMA, pierced by small, cribrate intercameral foramina. 1890)

Description: Agglutinated shell with subcy- Similarities and differences: The Pyrenean lindrical-to-subspherical morphology with multi- specimens are similar to those described by GEN- ple cribrate apertures piercing the apertural fa- DROT (1968) from the Martigues deposits. Howe- ce. The arrangement is planispiral-involute, but ver, the French specimens do not present grains may become uncoiled in the late stage of of quartz in their shells, suggesting that the growth. Pseudocyclammina presents exoskeleton presence of this type of agglutination in our composed of beams and rafters resulting in a specimens has an environmental significance. polygonal subepidermal network. It lacks endoskeleton. Specimens attributed to P. aff. sphaeroidea also occur in the Coniacian deposits of the Pyre- Age: After LOEBLICH & TAPPAN (1988), the age nees (CAUS et al., 2013). The specimens designed of Pseudocyclammina ranges from the Early as P. sphaeroidea in the Turonian of the Tethyan Jurassic to Coniacian, but the presence of P. deposits are smaller than the Pyrenean ones (see sphaeroidea in the Santonian (GENDROT, 1968) for instance, FRIJIA & PARENTE, 2008; CHIOCCHINI et and in the Early Campanian (this paper) indi- al., 2012; FRIJIA et al., 2007; ARRIAGA et al., in cates a younger age for this genus. press).

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Figure 12: Pseudocyclammina sphaeroidea. A. Oblique section showing three spiral whorls in a large specimen. B, E, H, K. Axial and subaxial sections. C-D, F-J. Equatorial and subequatorial sections. L. Centred oblique section. A, B, J from CO-30; C-E, H, I from CO-1; F, G, K, L from M-125. Scale bar 1 mm.

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Stratigraphical distribution: This small arranged, and not trochospirally arranged in the and globular Pseudocyclammina characterizes all early stages of growth as defined by LOEBLICH & deposits of the Font de les Bagasses Unit, TAPPAN (1964). Other characteristics, like a deep although it is most frequent in its lower part. At subepidermal network exoskeleton, the multiple certain levels it occurs as isolated specimens, apertures and the lack of endoskeleton are iden- but is more abundant in hard rocks. In the Pyre- tical to Dicyclina characteristics. Thus, in accor- nees, the species is known from Coniacian to dance with ARRIAGA et al. (in press), the genus Lower Campanian. However, it is also mentioned Cuneolina needs to be placed in the Family from the Turonian (FRIJIA & PARENTE, 2008; CHIO- Dicyclinidae. CCHINI et al., 2012; FRIJIA et al., 2007; among Therefore, the Family Dicyclinidae is emended others). here to include the non-cyclical forms as follows: Geographic distribution: P. sphaeroidea test free, conical to fan-shaped or discoidal, type is reported from the Pyrenean basin, while chamber biserially arranged, exoskeleton the small forms have a wider occurrence (see for composed of beams and rafters forming a regular instance: SCHLAGINTWEIT, 1992; FRIJIA et al., subepidermal network covered by a fine 2007). epidermis. Multiple apertures are placed at the Suborder Ataxophragmiina base of the chamber. It lacks endoskeletal structures. FURSENKO, 1958 Subfamily Cuneolininae SAIDOVA, 1981 Remarks: KAMINSKI (2014) included in this suborder "agglutinated high trochospiral forami- Remarks: ARRIAGA et al. (in press) maintained nifera with an asymmetrical spire that may re- the Subfamily Cuneolininae for the conical to fan- duce to a biserial or uniserial arrangement. The shaped morphotypes. However, further studies chambers may have exoskeletal and endoske- are needed to prove the validity of this subfamily, letal structures. The aperture forms a wide hori- comprising also small and less complex forms zontal slit at the base of the apertural face". from the Lower Cretaceous. However, some genera included in the suborder Cuneolina (ORBIGNY, 1839) have multiple apertures. ARNAUD-VANNEAU & SLITER, 1995 Superfamily Ataxophragmioidea (Type-species: Cuneolina pavonia ORBIGNY, SCHWAGER, 1877 1846) Remarks: After LOEBLICH & TAPPAN (1988), Description: Conical to fan-shaped for a- this Superfamily is characterized by its early minifera with agglutinated wall. The chambers trochospiral growth that may be reduced to are biserially arranged. They are generally broad biserial or uniserial in later stages. The cham- and low, and commonly compressed in the plan bers have skeletal elements partially dividing the of the biseriality. They have a complex exoske- chamber lumen. leton composed of beams and rafters forming a Family Dicyclinidae regular subepidermal polygonal network covered LOEBLICH & TAPPAN, 1964, emended by a fine epidermis. The beams frequently pene- trate deep into the chamber lumen leaving only a Remarks: ARRIAGA et al. (in press) narrow passage near the apertural face. It lacks considered the Family Dicyclinidae LOEBLICH & endoskeleton elements. Multiple intercameral TAPPAN, 1964, synonymous with Cuneolinidae foramina are aligned at the base of the septa. For SAIDOVA, 1981. LOEBLICH & TAPPAN (1964, 1978) more details see ARNAUD-VANNEAU & SLITER (1995). defined the Family Dicyclinidae as "test free, Age: Cuneolina is known from the Aptian to discoidal, chambers cyclical, biserially added, Maastrichtian (LOEBLICH & TAPPAN, 1988). partially subdivided by transverse or radial partitions with imperforate epidermal layer; Cuneolina conica ORBIGNY, 1850 peripheral multiple aperture". Whereas, the (Fig. 13 H, O ; Fig. 14 C-D) Family Cuneolinidae SAIDOVA, 1981, was descri- Synonymies: bed as "conical to subflabelliform, trochospiral in 1850 Cuneolina conica ORBIGNY, Pl. 1, figs. 7-9; Pl. 2, early stage, with up to five chambers per whorl, fig. 1. rapidly reduced to biserial; chambers subdivided 1899 Cuneolina conica ORBIGNY, SCHLUMBERGER, Pl. 8, by many septula extending from the outer wall figs. 8-10 toward the junction of the two series of cham- 1968 Cuneolina conica ORBIGNY, GENDROT, Pl. 4, fig. 16 bers¸ agglutinated wall with imperforate outer Description: Fan-shaped shell, biserially layer and reticulate subepidermal layer; aper- arranged, with low and broad chambers occu- ture simple to multiple". pying an opening angle of about 70º. The number According to SCHLAGINTWEIT (1992), ARNAUD- of chambers is about 20. The shell varies from VANNEAU & SLITER (1995) and the present work, 0.7mm to 1.2 mm in length. The exoskeleton Cuneolina ORBIGNY, 1839 (type-species: C. pavo- consists of well-developed beams and rafters. The nia ORBIGNY, 1846) shows a large and structured number of beams per chamber can reach about deuteroconch followed by chambers biserially 20 in the last chamber.

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Figure 13: A, C. Hemicyclammina cf. chalmasi: subaxial sections; from SG-8 and SG-10. B. oblique section not far from the equatorial section; from SG-8 and SG-10. D, E. Hemicyclammina cf. chalmasi:Oblique sections; from SG-8 and SG-10. F. Hemicyclammina cf. chalmasi: slightly oblique near the subaxial section; note in all of the pictures the coarse agglutinating quartz grains masking the wall structure; from SG-8 and SG-10. G, J, K, M. Cuneolina cyclindrica: axial sections; G, from M-130; K, from M-125; J, M from M-151. I. Cuneolina cyclindrica: slightly oblique section almost subaxial; from M-149. L. Cuneolina cyclindrica: basal section showing the beams penetrating deep into the chamber lumen and leaving a narrow passage; from M-130. N. Cuneolina cyclindrica: oblique section showing fifth chambers; from M-130. H, O. Cuneolina conica: longitudinal-oblique sections; respectively from M-143 and from M- 130. Scale bar 1 mm.

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Similarities and differences: Our speci- epidermis. They lack endoskeleton. mens from the Font de les Bagasses are identi- Age: The age attributed by LOEBLICH and TAP- cal to those described by SCHLUMBERGER (1899) PAN (1988) to this genus is Cenomanian-to- from Tragó de Noguera. Maastrichtian. Stratigraphical distribution: It is known Dictyopsella kiliani only from the Upper Cretaceous. MUNIER-CHALMAS in SCHLUMBERGER, 1899 Geographical distribution: Cuneolina coni- (Fig. 14 E-M ; Fig. 15 A-F) ca has been mentioned in the Pyrenean deposits. Synonymies: 1899 Dictyopsella kiliani MUNIER-CHALMAS in Cuneolina cylindrica HENSON, 1948 SCHLUMBERGER, Pl. 8, figs. 5, 7; Pl. 11, fig. 20. (Fig. 13 G, I-N ; Fig. 14 A-B) 1967 Dictyopsella kiliani MUNIER-CHALMAS in Synonymies: SCHLUMBERGER, HOTTINGER, Pl. 10, figs. 7-10. 1968 Dictyopsella kiliani MUNIER-CHALMAS in 1948 Cuneolina cylindrica HENSON, Pl. 14. fig. 7. SCHLUMBERGER, GENDROT, Pl. 5, figs. 7-10. 1968 Cuneolina cylindrica HENSON, GENDROT Pl. 4, fig. 1981 Dictyopsella kiliani MUNIER-CHALMAS in 14. SCHLUMBERGER, TRONCHETTI, Pl. 9, figs. 7-10; Pl. Description: Agglutinated, cylindrical, small 10, figs. 1-3 shell with biserially arranged chambers. The 1985 Dictyopsella kiliani MUNIER-CHALMAS in chambers grow regularly and have a subglobular SCHLUMBERGER, LOEBLICH & TAPPAN, Pl. 4, figs. 1-7; to slightly elongated morphology, opening with Fig. 3. an angle of 10º-15º (7º to 14º in HENSON, 1992 Dictyopsella kiliani MUNIER-CHALMAS in 1948). The shell varies from 0.5mm to 1.1 mm SCHLUMBERGER, SCHLAGINTWEIT, Figs. 7-10. in length. The wall consists of a fine epidermis Description: All the specimens found in the (recrystallized) and few developed beams and Font de les Bagasses unit are attributed to the rafters. A maximum of six beams per chamber type-species of the genus: D. kiliani. The main have been measured. The external wall and sep- characteristics of this species are reflected in the ta agglutinate coarse and angular quartz grains, generic description of SCHLUMBERGER (1899) from as occurs in many other foraminifera located in the deposits of the Tragó de Noguera. The shell the Les Bagasses deposits (see for instance, forms a low cone with its base plane or slightly Pseudocyclammina sphaeroidea). convex. The periphery is circular and slightly lo- bulated. The maximum diameter of the measured Stratigraphical distribution: This species specimens varies from 0.9 to 2.2 mm (A-forms). in known from the Santonian and Campanian In the dorsal side, they show 20-21 half-moon- deposits. shaped chambers arranged in two and half Geographical distribution Cuneolina cylin- whorls, with the last whorl having 8-10 cham- drica was described from the Middle East, but is bers. The dorsal sutures are arched and slightly a poorly known species. It is common in the depressed. In the ventral side, the chambers are Pyrenees. triangular and the sutures appear precisely at 2/3 Family Dictyopsellidae of its inner part of the shell and slightly curved at the 1/3 of the external margin. The embryo con- BRÖNNIMANN et al., 1983 sists of a spherical protoconch and a reniform to Remarks: Low trochospiral shell with poly- subglobular deuteroconch with exoskeleton. gonal subepidermal network. The adjacent Remarks: Although GENDROT (1968), in the chambers communicate by a main aperture in material from Martigues, attributed the small an interiomarginal position. specimens (with a diameter of about 1.3 mm) to Dictyopsella MUNIER-CHALMAS, 1899 the A-generation and the large ones (up to 2 mm) to the B-generation, all the measured speci- (Type-species: Dictyopsella kiliani MUNIER- mens from the Font de les Bagasses Unit are A- CHALMAS in SCHLUMBERGER 1899) forms Description: Finely agglutinated shell, with Similarities and differences: HOTTINGER low conical shape showing a convex dorsal side (1967), using material from Tragó de Noguera, and a plane or plano-convex umbilical side. The described the species D. muretae. This species, chambers are arranged in a low trochospire. In according to the author, differs from D. kiliani in the dorsal side, the chambers have the typical its larger number of chambers in the last whorl half-moon shape, while in the ventral side they (about 15) and the bigger size of the embryo are triangular leaving an open umbilicus in the compared with the embryo of D. kiliani. center of the shell. The main aperture is arched Moreover, HOTTINGER (1967) indicated that the and placed at the base of the apertural face. walls of the ventral side of the successive cham- Small supplementary openings are developed in bers are superposed, forming a columellar plate, the septal sutures. The interior of the chambers suggesting a higher degree of evolution for this is occupied by a deep exoskeleton, composed of last species. However, in the studied specimens numerous beams and rafters forming a fine collected from the Font de les Bagasses Unit, subepidermal polygonal network covered by the such characteristics have not been observed.

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Figure 14: A-B. Cuneolina cylindrica: external view of isolated specimens (scanning electron microphotographs); note the cylindrical morphology of this species. C-D. Cuneolina conica: lateral view of well preserved specimens. E-I. Dictyopsella kiliani: external view of the dorsal side showing the spiral half-moon shaped chambers (scanning microphotographies). J-L. Dictyopsella kiliani: ventral side with the chambers of the last whorl showing the main aperture and small septal openings. M-N. Details of J and L specimens. All from SG-36. Scale bar 1 mm.

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Figure 15: A-C. Dictyopsella kiliani from SG-36. A, C. traverse sections showing the disposition of chambers in the spiral side. B. traverse section showing the disposition of chambers in the last whorl.D. axial section of a small specimen showing the spherical protoconch and a reniform deuteroconch, from M-148. E- F. Subaxial sections, from M- 150. Scale bar 1 mm.

GENDROT (1968) described from Martigues, associated with porcellaneous and lamellar- under the name of D. cuvillieri, a small, thin and perforate foraminifera. Among the former group, flat species with circular outline, which was used lacazinids and meandropsinids are dominant, by LOEBLICH & TAPPAN (1985) to erect their new while the latter group is dominated by rotaliids genus Dictyopselloides. LOEBLICH & TAPPAN (1985) and orbitoidids. based the new genus on the presence of an The lacazinids are represented by two species exoskeleton formed only by beams or, if they that have replaced each other in time: Pseu- exist, a few rafters (see LOEBLICH & TAPPAN, 1988, dofabularia cantabrica HOTTINGER et al., 1989, and p. 150), while D. kiliani has well developed Lacazina elongata MUNIER-CHALMAS, 1899 (generic beams and rafters. However, the pictures and specific characteristics of these two taxa are presented by GENDROT (1968, Pl. 5, fig. 5) clearly treated in HOTTINGER et al., 1989). Meandropsinids indicate the presence of rafters; therefore, the are represented clearly by two small, apparently validity of the genus Dictyopselloides is nondimorphic forms: Nummofallotia cretacea questioned here. GENDROT (1968) differentiated (SCHLUMBERGER, 1899) and Eofallotia cf. simplex D. cuvillieri from D. kiliani by its smaller size, HOTTINGER & CAUS, 2009, which are present within flatter morphology, and the number of chambers complete unit. The large and extremely dimorphic in the last whorl, four to five. In our material, species Alexina payracea HOTTINGER & CAUS, 2009, morphotypes similar to those described by is frequent only in the lower part of the sections, GENDROT (1968) as D. cuvillieri occur only spora- whereas the larger B forms of Larrazetia larrazeti dically in thin-sections, not permitting a detailed (MUNIER-CHALMAS in SCHLUMBERGER, 1899) occur study. only in the upper part. The specimens designated Stratigraphical distribution: Coniacian- in the charts as Fascispira cf. schlumbergeri Lower Campanian. HOTTINGER & CAUS, 2009, need to be further Geographical distribution: Dictyopsella studied. Meandropsina vidali SCHLUMBERGER, 1899, kiliani is mentioned frequently from western and described from the facies of the Tragó de No- central Europe. guera, has been identified only in some washed samples collected in the middle part of the unit, 6. Comments on the associated and is omitted in the chart range (for more porcellaneous information see HOTTINGER & CAUS, 2009). and lamellar-perforate foraminifera Among the lamellar-perforate foraminifera, In the Font de les Bagasses Unit the the small Orbitoides hottingeri HINTE and O. san- agglutinated larger foraminifera are frequently tae-pelagiae ASTRE are frequent, and they replace

264 Carnets Geol. 15 (18) each other in time (ALBRICH et al., 2014). The Scientific Results, College Station, vol. 143, p. Family Rotaliidae (sensu HOTTINGER, 2014) is 537-564. represented by Rotorbinella spp., and Pyrenero- ARRIAGA M.E., FRIJIA, G., PARENTE M. & CAUS E. (in talia longifolia BOIX et al., 2009. The very small, press).- Benthic foraminifera in the aftermath heavily ornamented forms with areal foramina, of the Cenomanian-Turonian boundary extinc- umbilical flaps and tooth-plate are designated in tion event in the carbonate platform facies of this work as Pararotalia tuberculifera (REUSS, the southern Apennines (Italy).- Journal of 1861), but the ensemble should be further Foraminiferal Research, Lawrence. studied. The list of hyaline foraminifera ends BANNER F.T. (1970).- A synopsis of the Spiro- with the well-known Praestorsella roestae (VIS- cyclinidae.- Revista Española de Micropaleon- SER, 1951), Praesiderolites praevidali (NEUMANN, tologia, Madrid, vol. 2, nº 3, p. 243-290. 1987), and probably several species of the BOIX C., FRIJA G., VICEDO V., BERNAUS J.M., DI LUCIA genus Goupillaudina. M., PARENTE M. & CAUS E. (2011).- Larger foraminifera distribution and strontium isotope 7. Conclusions stratigraphy of the La Cova limestones The detailed architectural study of the fora- (Coniacian - Santonian, Serra del Montsec, minifera collected in the Font de les Bagasses Pyrenees, NE Spain).- Cretaceous Research, Unit, in the Montsec Mountains, has allowed the vol. 32, p. 806-822. following agglutinated foraminifera to be BOIX C., VILLALONGA R., CAUS E. & HOTTINGER L. identified: Choffatella aff. rugoretis, Hemicy- (2009).- Late Cretaceous rotaliids (Foramini- clammina chalmasi, Pseudocyclammina sphae- ferida) from the Western Tethys.- Neues roidea, Cuneolina conica, C. cylindrica and Jahrbuch für Geologie und Paläontologie, Dictyopsella kiliani. All of these share the same Stuttgart, vol. 253, nº 2-3, p. 197-227. type of exoskeleton composed of beams and CAUS E. (1981).- Structural analysis of larger rafters with a fine epidermis (polygonal subepi- foraminifera in random section as an instru- dermal network) that is combined with different ment for rapid determination of rock age and arrangements of chambers to conform: environment. In: MARTINELL J. (ed.), Interna- 1. Planispiral to uncoiled, peneropliform or tional Symposium on Concept and Method in cyclical (Spirocyclinidae); Palaeontology.- Departament de Paleontolo- 2. Biserial (Dicyclinidae); gia, Universitat de Barcelona, p. 223-232. 3. Low trochospiral (Dictyopsellidae). CAUS E. & CORNELLA A. (1983).- Macroforami- In the suprageneric classification Choffatella nifères du Crétacé supérieur du Bassin Sud- is included in Choffatelinae MAYNC, Hemicyclam- Pyrénéen.- Géologie Méditerranéenne, Mar- mina in Hemicyclammininae BANNER, and Pseu- seille, vol. 13, nº 4, p. 137-142. docyclammina in Amijiellinae SEPTFONTAINE within CAUS E., CORNELLA A. & PONS J.M. (1978).- Forami- the Spirocyclinidae. Cuneolina is included in the níferos bentónicos del Santoniense surpi- Family Dicyclinidae. renaico (Montsec de Rubies, prov. de Lerida, España). Nueva adscripción genérica de Dic- Acknowledgements tyopsella chalmasi SCHLUMBERGER.- Revista We wish to express our sincere gratitude to Española de Micropaleontología, Madrid, vol. Prof. Joan ROSELL for his help in the field work. 10, nº 3, p. 453-460. Particular thanks are due to the reviewers Dr. CAUS E. & GÓMEZ-GARRIDO A. (1989a).- Correlation Annie ARNAUD-VANNEAU, Dr. Felix SCHLAGINTWEIT of larger benthic and planktonic foraminifera and to Prof. Bruno GRANIER (editor) whose of the Late Cretaceous in the South-Central comments considerably improved this work. This Pyrenees. In: WIEDMAN J. (ed.), Cretaceous of research was funded by the Spanish Ministry of the western Tethys. Proceedings of the 3rd Science and Innovation (project CGL2009- International Cretaceous Symposium 08371) and Economy and Competitivity (project (Tübingen).- E. Schweizerbart'sche Verlags- CGL2012-33160). bucchandlung, Stuttgart, p. 231-238. CAUS E. & GÓMEZ-GARRIDO A. (1989b).- Upper Bibliographic references Cretaceous bioestratigraphy of the south- ALBRICH S., FRIJIA G., PARENTe M. & CAUS E. central Pyrenees (Lleida, Spain).- Geodinamica (2014).- The evolution of the earliest repre- Acta, Paris, (Series 3), vol. 3, p. 221-228. sentatives of the genus Orbitoides: impli- CAUS E., LLOMPART C., ROSELL J. & BERNAUS J.M. cations for Upper Cretaceous biostrati- (1999).- El Coniaciense superior-Santoniense graphy.- Cretaceous Research, vol. 51, p. 22- inferior de la Sierra del Montsec (Pirineos, NE 34. de España).- Revista de la Sociedad Geológica ARNAUD-VANNEAU A. & SLITER W.V. (1995).- Early de España, Salamanca, vol. 12, nº 2, p. 269- Cretaceous shallow-water benthic forami- 280. nifers and fecal pellets from leg 143 CAUS E., PARENTE M., VICEDO V., FRIJIA G. & MAR- compared with coeval faunas from the Pacific TINEZ R. (2013).- Broeckina gassoensis sp. basin, Central America, and the Tethys.- nov., a larger foraminiferal index fossil for the Proceeding of the Ocean Drilling Program, middle Coniacian shallow-water deposits of the Pyrenean Basin (NE Spain).- Cretaceous Re-

265 Carnets Geol. 15 (18)

search, vol. 45, p. 76-90. HINTE J.E. van. (1966).- Orbitoides hottingeri n. CHIOCCHINI M., PAMPALONI M.L. & PICHEZZI R.M. sp. from Northern Spain.- Proceedings of the (2012).- Microfacies e microfossili delle suc- Koninklijke nederlandse Akademie van W- cessioni carbonatiche mesozoiche del Lazio e etenschappen, Amsterdam, (Series B), vol. dell'Abruzzo (Italia centrale) - Cretacico.- Me- 69, nº 3, p. 388-402. morie per servire alla descrizione della Carta HOTTINGER L. (1967).- Foraminifères imperforés du Geologica d'Italia, Roma, p. 269. Mésozoïque marocain.- Notes et Mémoires du CLAPÉS S. (1997, unpublished).- Els nivells de Service géologique, Rabat, vol. 209, p. 5-168. gresos santonians de les Serres Marginals HOTTINGER L. (1978).- Comparative anatomy of (Prepirineu català).- MSc Thesis, Universitat elementary shell structure in selected larger Autònoma de Barcelona, Bellaterra, 151 p. Foraminifera. In: HEDLEY R.H. & ADAMS C.G. [in Catalan]. (eds.), Foraminifera.- Academic Press, Lon- CORNELLA A. (1977).- Foraminíferos bentónicos don, vol. 3, p. 204-266. del Santoniense del Barranco de la Font de la HOTTINGER L. (2006).- Illustrated glossary of Plata, Montsec de Rúbies, prov. de Lérida.- terms used in foraminiferal research.- Carnets Publicaciones de Geología de la Universidad Geol., Madrid, vol. 6, nº M02 (CG2006_M02), Autónoma de Barcelona, Bellaterra, nº 8, 45 p. 125 p. DERCOURT J., ZONENSHAIN L.P., RICOU L.E., KAZMIN HOTTINGER L. (2014).- Paleogene larger Rotaliid V.G., LE PICHON X., KNIPPER A.L., GRANDJACQUET Foraminifera from the Western and Central C., SBORSHCHIKOV J.M., BOULIN J., SOROKHTIN Neothetys.- Springer, p. 3-191. O., GEYSSANT J., LEPVRIER C., BIJU DUVAL B., HOTTINGER L. & CAUS E. (2009).- Meandropsinidae, SIBUET J.C., SAVOSTIN L.A., WESTPHAL M. & an ophtalmidid family of Late Cretaceous K- LAUER J.P. (1985).- Présentation de 9 cartes strategist foraminifera endemic in the paléogéographiques au 1/20 000 000, s'éten- Pyrenean Gulf.- Neues Jahrbuch für Geologie dant de l'Atlantique au Pamir pour la période und Paläontologie, Stuttgart, vol. 253, nº 2-3, du Lias à l'Actuel.- Bulletin de la Société géo- p. 249-279. e logique de France, Paris, (8 Série), t. I, nº 5, HOTTINGER L., DROBNE K. & CAUS E. (1989).- Late p. 637-652. Cretaceous, Larger, Complex Miliolids (For a- DOUVILLÉ H. (1906).- Évolution et enchaînement minifera) endemic in the Pyrenean Faunal des Foraminifères.- Bulletin de la Société Province.- Facies, Erlangen, vol. 21, p. 99- géologique de France, Paris, (4e Série), t. IV, 134. p. 588-602 (Pl. XVIII). KAMINSKI M.A. (2004).- The year 2000 classi- FRIJIA G. & PARENTE M. (2008).- Reticulinella fication of the agglutinated foraminifera. In: kaeveri CHERCHI, RADOIČIĆ and SCHROEDER: a BUBIK M. & KAMINSKI M.A. (eds.), Proceedings marker for the middle upper Turonian in the of the fifth international workshop on agglu- shallow-water carbonate facies of the peri- tinated Foraminifera.- Grzybowski Foundation adriatic area.- Bollettino della Società Geolo- Special Publication, Krakow, vol. 8, p. 237- gica Italiana, Roma, vol. 127, p. 275–284. 255. FRIJIA G., PARENTE M., DI LUCIA M. & MUTTI M. KAMINSKI M.A. (2014).- The year 2010 classi- (2007).- Carbon and strontium isotope fication of the agglutinated foraminifera.- stratigraphy of the Upper Cretaceous (Ceno- Micropaleontology, New York, vol. 60, nº 1, p. manian-Campanian) shallow-water carbo- 89-108. nates of southern Italy: Chronostratigraphic LOEBLICH A.R. & TAPPAN H. (1964).- Sarcodina, calibration of larger foraminifera biostrati- chiefly "Thecameobians" and Foraminiferida. graphy.- Cretaceous Research, vol. 53, p. In: MOORE R.C. (ed.), Treatise on Invertebrate 110-139. Palaeontology, Part C, Protist 2.- Geological GENDROT C. (1968).- Stratigraphie et micropa- Society of America, Boulder, and University of léontologie du Sénonien de la région des Mar- Kansas, Lawrence, 900 p. tigues près Marseille (Bouches-du-Rhône).- LOEBLICH A.R. & TAPPAN H. (1985).- Some new and Eclogae Geologicae Helvetiae, Basel, vol. 61, redefined genera and families of agglutinated nº 2, p. 657-705. foraminifera; I.- Journal of Foraminiferal GRADSTEIN F.M., OGG J.G., SCHMITZ K. & OGG G. Research, Lawrence, vol. 15, p. 91-104. (2012).- The Geologic Time Scale 2012.- LOEBLICH A.R. & TAPPAN H. (1988).- Foraminiferal Elsevier, 2 volumes, 1144 p. genera and their classification.- Van Nostrand HAY W.W., DE CONTO R.M., WOLD C.N., WILSON Reinhold, New York, 970p. K.M., VOIGHT S., SCHULZ M., WOLD A.R., DULLO LOEBLICH A.R. & TAPPAN H. (1992).- Present status W.C., RONOV A.B., BALUKHOVSKY A.N. & SÖDING of Foraminiferal classification. In: TAKAYANAGI E. (1999).- Alternative global Cretaceous Y. & SAITO T. (eds.), Studies in benthic Fora- paleogeography.- Geological Society of minifera.- Tokai University, Tokyo, p. 93-102. America, Special Paper, Washington D.C., nº LOEBLICH A.R. & TAPPAN H. (1994).- Foraminifera of 332, 47 p. the Sahul Shelf and Timor Sea.- Cushman HENSON F.R.S. (1948).- Larger imperforate For a- Foundation for Foraminiferal Research, Special minifera of South-Western Asia.- British Publication, Lawrence, vol. 31, 661 p. Museum (Natural History), London, 127 p. MAYNC W. (1950).- The foraminiferal genus

266 Carnets Geol. 15 (18)

Choffatella SCHLUMBERGER in the Lower Creta- composition of fish tooth enamels.- Paleo- ceous (Urgonian) of the Caribbean region ceanography, Washington D.C., vol. 18, nº 2, (Venezuela, Cuba, Mexico and Florida).- Eclo- p. 1-12. gae geologicae Helvetiae, Basel, vol. 42, nº RADOIČIĆ R., RADULOVIĆ V., RABRENOBIĆ D. & RADU- 2, p. 529-547. LOVIĆ B. (2010).- The age of the brachiopod MAYNC W. (1953).- Hemicyclamina sigali, n. gen. limestones from Guča, western Serbia.- Anna- nov. sp. from Cenomanian of Algeria.- Con- les géologiques de la Péninsule Balkanique, tributions from the Cushman Foundation for vol. 71, p. 73-93. Foraminiferal Research, Lawrence, vol. 4, nº SCHLAGINTWEIT F. (1992).- Benthonische Forami- 9, p. 148-149. niferen aus Flachwasserkarbonaten der Ober- MAYNC W. (1958).- Feurtillia frequens, n. gen., n. kreide der Nördlichen Kalkalpen (Gosau- sp., a new genus of Lituolid foraminifera.- schichtgruppe, Österreich).- Mitteilungen der Contributions from the Cushman Foundation Österreichischen geologischen Gesellschaft, for Foraminiferal Research, Lawrence, vol. 9, Wien, vol. 84, p. 327-353. part. 1, p. 1-3. SCHLAGINTWEIT F. & WAGREICH M. (2004).- Hemi- MAYNC W. (1972).- Gendrotella n. gen. and Chof- cyclammina chalmasi (SCHLUMBERGER, 1899) - fatella caronae n. sp. from the Lower Seno- Larger benthic Foraminifera from the Upper nian of Southern France.- Eclogae geologicae Cretaceous Gosau Group of the Northern Helvetiae, Basel, vol. 65, p. 355-359. Calcareous Alps (Gosau, Austria).- Jahrbuch MILLER K.G., KOMINZ M.A., BROWNING J.V., WRIGHT der geologischen Bundesanstalt, Wien, vol. J.D., MOUNTAIN G.S., KATZ M.E., SUGARMAN 144, p. 367-372. P.J., CRAMER B.S., CHRISTIE-BLICK N. & PEKAR SCHLUMBERGER C. (1899).- Note sur quelques S.F. (2005).- The Phanerozoic Record of foraminifères nouveaux ou peu connus du Global Sea-Level Change.- Science, Wa- Crétacé d'Espagne.- Bulletin de la Société shington D.C., vol. 310, p. 1293-1298. géologique de France, Paris, (3e Série), t. MUNIER-CHALMAS E. (1887).- Sur la Cyclolina et XXVII, p. 456-465 (Pls. VIII-XI). trois nouveaux genres de foraminifères des SEPTFONTAINE M. (1980).- Les Foraminifères imper- couches à rudistes: Cyclopsina, Dicyclina et forés des milieux de plate-forme au Méso- Spirocyclina.- Compte Rendu des Séances de zoïque : détermination pratique, interprétation la Société géologique de France, Paris, vol. 4, phylogénétique et utilisation biostratigra- nº 7, p. 30-31. phique.- Revue de Micropaléontologie, Paris, MUÑOZ J.A. (1992).- Evolution of a continental vol. 23, nº 3-4, p. 169-203. collision belt: ECORS-Pyrenees crustal balan- SEPTFONTAINE M. (1988).- Vers une classification ced cross-section. In: MCCLAY K.R. (ed.), évolutive des Lituolidés (Foraminifères) juras- Thrust tectonics.- Chapman & Hall, London, siques en milieu de plate-forme carbonatée.- p. 235-246. Revue de Paléobiologie, Genève, vol. spec. nº NOGUÉS A. (2000, unpublished).- Biostromas de 2 (Benthos '86), p. 229-256. corales, rudistas y esponjas coralinas del SIMÓ A. (1986).- Carbonate platform depositional Santoniense del Montsec d'Ares: carac- sequences, Upper Cretaceous, south-central terización estratigràfica, sedimentológica y Pyrenees (Spain).- Tectonophysics, vol. 129, paleontológica.- MSc Thesis, Universitat p. 205-231. Autónoma de Barcelona, Bellaterra, 216 p. TEIXELL A. & MUÑOZ J.A. (2000).- Evolución tecto- ORBIGNY A. d' (1850).- Prodrome de Paléontolo- nosedimentaria del Pirineo meridional durante gie stratigraphique universelle des animaux el Terciario: una síntesis basada en la mollusques et rayonnés.- Masson, Paris, vol. transversal del río Noguera- Ribagorçana.- 2, 428 p. Revista de la Sociedad geológica de España, PAWLOWSKI J., HOLZMANN M. & TYSZKA J. (2013).- Salamanca, vol. 13, nº 2, p. 251-264. New supraordinal classification of Forami- TRONCHETTI G. (1981).- Les foraminifères crétacés nifera: Molecules meet morphology.- Marine de Provence (Aptien-Santonien). Systéma- Micropaleontology, vol. 100, p. 1-10. tique, biostratigraphie, paléoécologie, paléo- POLAVDER S. (1999).- On some little known géographie.- PhD Thesis, Université de Pro- foraminifers from the Senonian of the Vardar vence ; Travaux du Laboratoire de Géologie Zone.- Acta Palaeontology Romaniae, Cluj- Historique et de Paléontologie, Marseille, vol. Napoca, vol. 2, p. 373-375. 12, nº 3, 559 p. PONS J.M. (1977).- Estudio estratigráfico y VILLALONGA R. (2009, unpublished).- Los macrofo- paleontológico de los yacimientos de Rudistas raminíferos de la plataforma Campaniense del del Cretácico Superior del Prepirineo de la margen sur de la cuenca pirenaica (Unidad provincia de Lérida.- Publicaciones de Geolo- Surpirenaica central, NE de la Península Ibé- gía, Universitat Autònoma de Barcelona, rica).- PhD Thesis, Universitat Autònoma de Bellaterra, nº 3, 105 p. Barcelona, Bellaterra, 216 p. PUCÉAT E., LÉCUYER C., SHEPPARD S.M.F., DROMART WANNIER M. (1983).- Évolution, biostratigraphie et G., REBOULET S. & GRANDJEAN P. (2003).- systématique des Siderolitinae (Foraminifè- Thermal evolution of Cretaceous Tethyan res).- Revista Española de Micropaleontologia, marine waters inferred from oxygen isotope Madrid, vol. 15, nº 1, p. 5-37.

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