Cuvillierinella salentina (Foraminifera, Rhapydioninidae) and its kinship in the Western Mediterranean area during the Campanian–Maastrichtian

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Original article

Cuvillierinella salentina (Foraminifera, Rhapydioninidae) and its kinship in the Western Mediterranean area during the Campanian–Maastrichtian Cuvillierinella salentina (Foraminifères, Rhapydioninidae) et ses proches en Méditerranée occidentale au Campanien–Maastrichtien Jean-Jacques Fleury 118, avenue de Flandre, 59290 Wasquehal, France

Abstract Cuvillierinella salentina Papetti and Tedeschi, 1965 and its related species of the Rhapydioninidae family are widely distributed in the Western Mediterranean region by Campanian–Maastrichtian time. C. salentina, the type species of the genus, is studied in rich populations from its Italian type locality, and several other spots from Greece and Spain. It shows great variability: the well-known type coexists with streptospiral tests almost devoid of endoskeleton and wholly planispiral tests with fine mesh-like endoskeleton, as well as many intermediates. Such populations reflect the large capacity of evolution of at least the group of miliolids from which these populations are derived, if not the species itself. The genera Murciella and Cyclopseudedomia are probably the most direct offshoots, but other taxa appear to arise from this species or at least from the same stock of origin. In the same genus and at the same time (middle part of Campanian age, “CsB6a” zone), two new species are described, both hesitating between the streptospiral and planispiral coiling: – C. fluctuans nov. sp., from Greece, A tests being either streptospiral or planispiral, with primary and secondary chamberlets taking the aspect of polygonal isodiametric network and – C. perisalentina nov. sp., from Italy, with persistent streptospiral coiling and disordered arrangement of secondary chamberlets which allow us to consider its relation with species of the genus Pseudochubbina, hitherto of completely enigmatic origin. Assigned to the same genus, from Upper Campanian–Lower Maastrichtian time (“CsB6b” zone), C. aff. pylosensis, probably related to C. pylosensis, is studied from several populations, and constitutes, with the above mentioned species, what is called C. gr. salentina. Based on material from the CsB6b zone, the new genus Metacuvillierinella nov. gen. is introduced; M. decastroi nov. sp., type species of the new genus, known from Greece and Italy, shows characters common to C. gr. salentina, such as milioline to streptospiral nepionic coiling, large endoskeleton mesh, associated with some original characters, such as the unusual conjunction of the advolute coiling and absence of any final unrolling together with very low dimorphism of generation; this results in flat tests, often sigmoid shaped in axial section, very rare in the family. Thus, the genus Cuvillierinella, and especially C. salentina, appears as a possible source, or at least not far from the origin, of a number of taxa related to one another, constituting a large part of the Rhapydioninidae family. They are gathered together inside the new subfamily Cuvillierinellinae, distinct from the subfamily Rhapydionininae sensu stricto (comprising Rhapydionina and Fanrhapydionina) which makes a different and parallel branch, from Upper Campanian (CsB6b zone) to the end of Cretaceous time (CsB7 zone). © 2016 Published by Elsevier Masson SAS. Keywords: Foraminifera; Alveolinacea; Rhapydioninidae; Cuvillierinella salentina; Uppermost Cretaceous (Campanian–Maastrichtian)

Résumé Appartenant à la famille des Rhapydioninidae, Cuvillierinella salentina et les espèces apparentées sont largement répandues au Campanien–Maastrichtien dans la région méditerranéenne occidentale. C. salentina, l’espèce type du genre, est examinée à partir de riches populations provenant de sa localité type italienne et de plusieurs localités de Grèce et d’Espagne. On montre sa grande variabilité: formes streptospiralées presque dépourvues d’endosquelette et formes entièrement planispiralées à maille endosquelettique fine, outre de nombreux intermédiaires, coexistent avec le type bien connu. De telles populations témoignent de grandes capacités d’évolution, sinon de l’espèce même, du moins du groupe de Miliolidés dont ces populations sont issues. Les genres Murciella et Cyclopseudedomia en sont probablement les plus directs descendants, mais d’autres taxons en paraissent découler ou provenir du même stock d’origine. Dans le même genre et à la même époque E-mail address: [email protected] http://dx.doi.org/10.1016/j.revmic.2016.04.002 0035-1598/© 2016 Published by Elsevier Masson SAS.

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(partie moyenne du Campanien, zone «CsB6a»), sont décrites deux espèces nouvelles, hésitant entre les enroulements streptospiralé et planispiralé: C. fluctuans nov. sp., de Grèce, dont les tests A sont soit streptospiralés, soit planispiralés, à logettes primaires et secondaires prenant un aspect de réseau à mailles polygonales isodiamétriques et C. perisalentina nov. sp. d’Italie, dont l’enroulement streptospiralé persistant et la disposition désordonnée des logettes secondaires permettent de suspecter des relations avec les espèces du genre Pseudochubbina, d’origine jusqu’à présent entièrement énigmatique. Dans le même genre encore, mais plus tardive (partie supérieure du Campanien et partie basale du Maastrichtien, zone «CsB6b»), C. aff. pylosensis proche mais mieux caractérisée que C. pylosensis est étudiée à partir de plusieurs populations. L’ensemble des espèces précédemment citées correspond à l’entité nommée Cuvillierinella gr. salentina. De la zone «CsB6b», est créé le nouveau genre Metacuvillierinella nov. gen. M. decastroi nov. sp., l’espèce type du nouveau genre, connue de Grèce et d’Italie, associe des caractères communs aux espèces de C. gr. salentina, tels que l’enroulement népionique streptospiralé à miliolin et l’endosquelette à large maille, à des caractères originaux, tels que l’insolite concomitance de l’enroulement advolute à l’absence de tout déroulement final et le très faible dimorphisme de générations. Une telle association de caractères n’est connue chez aucune des populations plus anciennes ou contemporaines de la famille; il en résulte des tests très plats, souvent d’allure sigmoïde en section axiale, également sans équivalent dans la famille. Ainsi, C. salentina apparaît comme à l’origine, ou proche de l’origine, de tout un ensemble de taxons clairement apparentés entre eux constituant une grande partie de la famille Rhapydioninidae; on les réunit au sein de la nouvelle sous famille Cuvillierinellinae, formant une branche parallèle à celle de la sous famille Rhapydionininae sensu stricto (réunissant Rhapydionina et Fanrhapydionina) du Campanien supérieur (zone CsB6b) à la fin du Crétacé (zone CsB7). © 2016 Publi´e par Elsevier Masson SAS. Mots clés : Foraminifères ; Alveolinacea ; Rhapydioninidae ; Cuvillierinella salentina ; Crétacé terminal (Campanien–Maastrichtien)

1. Introduction This work is the second of the “memoirs” of a geologist who, during his career, has had the opportunity to meet, fortuitously at first, then intentionally, many representatives of the family Rhapydioninidae, part of the super family Alveolinacea. This family was almost unknown in the mid 1960s and its rare identified members were not properly understood at that time. This discretion appears linked to very narrow ecological requirements, confining the group in restricted circulation platform environments, poorly represented in the fossil record. The counterpart of this confinement is that in favorable environments, protected from competition, populations grow abundantly and provide exceptional opportunities to study. A few morphologically distinct species, represented by several populations in a somewhat broad area lend themselves to confrontations from which emerge the immutable and more versatile characters submitted to ecology and evolution. Such an attempt was previously outlined (Fleury, 2014) about the relatives of Rhapydionina liburnica (Stache, 1889) and will be developed below for the species of the genus Cuvillierinella Papetti and Tedeschi, 1965 and its affiliate Metacuvillierinella nov. gen. 2. Abbreviations In order to summarize the uncertainties of attribution, or the relationships between taxa, the following formulas are used: “aff.” reports on probable differences with the named species, although affinities between them cannot be suspected, “spp.” (in the figures) refers to several species belonging to the same genus, without risk of ambiguity, “gr.” corresponds to a group of species of the same genus, of which other independent species are also known. With reference to the reproductive cycle of foraminifera, as it is usual, tests of megalospheric generation are called “A” and tests of microspheric generation are called “B”. Some

abbreviations are unique to this work. Biozones related to confined environments of the Adriatico-Aegean platforms are baptized “CsB”, and they are illustrated below. Furthermore, some recurrent composed terms are used in the text and the figures: UUT means Uncoiled Uniserial Termination (pseudoevolute or evolute, cylindrical or flaring), the final stage of many tests; SAR means Slow Axial Rotation, a special case of streptospiral coiling, illustrated particularly by species of the genera Pseudochubbina and Metacuvillierinella nov. gen.; SSC means Scattered Secondary Chamberlets, describing the disordered appearance of chamberlets imbedded in the “central endoskeleton” when well differentiated “floors” are missing (see Section 5). 3. Materials and methods The two genera that are the subject of this study are mainly prevalent in the internal areas of carbonate platforms in the vicinity of the Adriatic, more rarely in the Eastern Mediterranean, approximately as Rhapydionina (see Fleury, 2014, fig. 2) and also in Spain (Province of Murcia). The studied material is part of collections gathered by the author during a period of about twenty years, on the occasion of field expeditions funded by the University of Lille 1 and several formations associated with the French Centre National de la Recherche Scientifique (CNRS). It is made up of populations chosen as to be the most significant among those encountered mainly in mainland Greece and Italy. In addition, some studied samples are donations or loans due to several colleagues whose names will be mentioned on the occasion. The whole material is composed of thin sections carved from disaggregation-resistant limestones. These slices are brought together in a collection, where they are identified by a sample number preceded by a group of 3 letters peculiar to various locations in Greece (see Fig. 1), or the letter I for Italy. A collection number, also inscribed on each glass blade, preceded by the initials of the author (JJF), can be found either in the summary table of

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Fig. 1. Localization of some Rhapydioninidae bearing outcrops, from Greece and Italy. On the map, in Greece, the groups of three letters indicate the location of several outcrops from which the quoted samples of the text are numbered. XGP: Gavrovo massif (Valtou Ori); GKL: Klokova massif; GDD: Mainalon massif (near Vitina); GGB: Pylos-Methoni area; FPM: Astypalya island. In Italy, samples I250 (locality of Metacuvillierinella decastroi nov. gen., nov. sp. and Fanrhapydionina aff. flabelliformis of Fleury, 2014) and I340 (type locality of Cuvillierinella salentina, C. perisalentina nov. sp. and Pseudochubbina bruni) are from the Salentina peninsula. To the right, the Klokova section (GKL) displays the only continuous series of Rhapydioninidae bearing outcrops in Greece, from which several species are mentioned in the text. Abbreviations: Cuv.: Cuvillierinella; Cycl.: Cyclopseudedomia; Rhap.: Rhapydionina.

Fig. 12, either in the course of descriptions when absent in this figure. This collection is housed in the public paleontological collections of the University of Lille, 1. Sciences and Technology. The used analytical methods are very common, and they mainly consist in the study of thin sections of cemented rocks. Nevertheless, they differ from those that are ordinarily used by confrontation of different populations supposedly belonging to the same taxon. It is therefore interesting to identify, in a geographical area as wide as possible, changes in features which, beyond their diversity, characterize the studied taxon: it is obviously essential to define types, but they are far from being sufficient for the knowledge of a taxon. Moreover, as random sections given by the usual manufactured thin sections are sometimes insufficient for careful work, oriented and fitted sections were specially prepared in order to know the morphology of rare tests, such as those of B generation, for example. The main criteria of distinction of the species and genera are as follows. The internal diameter of proloculus, primordial element of the characterization of each population is always available and usually discriminating in its statistical aspect.

The coiling mode, variable during ontogenesis and sometimes difficult to characterize in random section, is also important; it constitutes the essential distinctive criterion for the genus Cuvillierinella. In that genus, in a general way, the first turns of whorl around the proloculus appear “irregular” in both generations, at least in the ordinary random sections at hand. The following terms are used to characterize its organization: streptospiral is the most comprehensive, meaning that the chamber arrangement does not seem to follow any simple organized system; milioline means that one can distinguish a system related to that of certain Miliolacea, in which the chambers (at least the very first ones) seem to be organized according to several planes convergent on the proloculus, such as in the system known as “pseudotriloculin” (Pêcheux, 2002); quinqueloculine means that one can distinguish an arrangement of chambers according to five planes, of the type which characterizes the genus Quinqueloculina d’Orb., 1826, assumed to be common to all the primitive stocks of Alveolinacea. The chamber partitioning (“endoskeleton”) is independent from the two previous criteria; the stage of its advent is always to be noted but it does not appear decisive for characterization of species and even populations. The appearance of the partitioning of the adult chambers, either likely consisting

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in layers of tubular chamberlets (separated by the so-called “floors”), either consisting in tubular chamberlets scattered in a compact mass (SSC), is a former subject of reflection to which it will be added a few new elements. The final unrolling of A test, remarkable but never a majority, appears ordinarily insignificant; however, its absence, if confirmed, can be characteristic (Metacuvillierinella nov. gen. for example). The B tests, potentially very significant, being more conservative in their nepionic part and more advanced in the adult, are not to be neglected, but they are often too scarce (sometimes absent) to allow reliable conclusions.

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4. Biostratigraphy The studies of Fleury (1980, 2014), Mavrikas (1993) and Landrein (2001) have led to the establishment of a biozonation of Upper Cretaceous endemic organisms from carbonates of the internal Gavrovo-Tripolitza platform in Greece. Its broad outlines are recognized in Dalmatia (Pejovi´c and Radoiˇci´c, 1987; Guˇsi´c and Jelaska, 1990), South Italy (Luperto Sinni and Ricchetti, 1978; Reina and Luperto Sinni, 1993) and Albania (Heba and Prichonnet, 2009). The biozones which concern this study correspond roughly to the Santonian–Maastrichtian period (Fig. 2).

Fig. 2. Campanian–Maastrichtian biostratigraphic zonation of the Greek Gavrovo-Tripolitza platform, after Fleury (1980, 2014), Mavrikas (1993) and Landrein (2001). The biozones (“CsB4 to CsB7”), defined by the distribution of endemic foraminifera (from “Restricted internal platform”), are tentatively correlated to the chronostratigraphic scale through the occasionally intercalated “Open shelf” foraminifera. The uncertainty of correlations is expressed by dotted lines on both sides of the “Open shelf” column. To the right, a “Sedimentation – diagenesis” column expresses schematically the environmental conditions: white corresponds to subtidal sediments, gray indicates inter- to supratidal facies and diagenesis. The continuous gray line between CsB6 and CsB7 illustrates a generalized subaerial exposure interpreted as responsible for the radical change in the populations. The thin discontinuous line between CsB6a and CsB6b illustrates the hypothesis of such an event at this time. Abbreviations: O.: Orbitoides; P.: Pseudosiderolites; K.: Keramosphaerina; M.: Moncharmontia. Pseudochubbina spp. corresponds to P. bruni De Castro, 1990 and P. philippsoni (Fleury, 1977); Murciella gr. renzi corresponds to M. renzi Fleury, 1979a and M. ovoidea Fleury, 1979a. Cyclopseudedomia spp. corresponds to C. smouti Fleury, 1974, C. klokovaensis (Fleury, 1979a), and C. hellenica Fleury, 1979b. Rhapydionina gr. dercourti includes R. dercourti Fleury, 2014, R. fourcadei Fleury, 2014, and R. fleuryi Vicedo and Caus, 2011 (of which R. “bulbiformis” of Fleury, 2014 is a junior synonym). Rhapydionina aff. liburnica is the Greek variety of Stache’s species (with relatively small proloculus: see Fleury, 2014). Unspecified authors of species are named in the text.

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The “CsB4” zone corresponds to the maximum extension of Murgella lata Luperto-Sinni, 1966 and, at least in Greece, to the presence of Keramosphaerina tergestina (Stache, 1889). The “CsB5” zone is not precisely characterized by sporadic Moncharmontia apenninica (De Castro, 1966), and Reticulinella fleuryi Cvetko, Guˇsi´c and Schroeder, 1997, already present in the previous zone. According to Cvetko et al. (2001), K. tergestina could accompany these species until the top of the zone. Senalveolina aubouini Fleury, 1984 is very rare in Greece but is present in Albania (author’s unpublished observation). The “CsB6” zone was originally (Fleury, 1980) defined by the presence of the Rhapydioninidae (excluding Rhapydionina gr. liburnica); two successive associations can be distinguished, after observations of Mavrikas (1993) in the Gavrovo massif (XGP, Fig. 1). A “CsB6a” association, ordinarily made up of Cuvillierinella salentina Papetti and Tedeschi, 1965 and Murciella cuvillieri Fourcade, 1966, to which is added Fleuryana adriatica De Castro, Drobne and Guˇsi´c, 1994. Other rarer species have to be mentioned: Pseudochubbina bruni De Castro, 1990 in Italy, P. philippsoni (Fleury, 1977) in Greece, as well as the two new species of the genus Cuvillierinella described in this work. A “CsB6b” association brings together, in various combinations, Murciella renzi Fleury, 1979a, M. ovoidea Fleury, 1979b, Cyclopseudedomia smouti Fleury, 1974, C. klokovaensis (Fleury, 1979b), C. hellenica Fleury, 1979b, Cuvillierinella pylosensis in Vicedo and Caus, 2011, C. aff. pylosensis (this work), Rhapydionina dercourti Fleury, 2014, R. fourcadei Fleury, 2014, R. fleuryi in Vicedo and Caus, 2011 (of which R. “bulbiformis” in Fleury, 2014 is a younger synonym), Metacuvillierinella perisalentina nov. gen., nov. sp., and Fanrhapydionina aff. flabelliformis (in Fleury, 2014). The “CsB7” zone is characterized by Rhapydionina gr. liburnica of which Greek representatives are perhaps not conspecific with R. liburnica (Stache) (see R. aff. liburnica in Fleury, 2014), associated with Neobalkhania bignoti Cherchi, Radoiˇci´c and Schroeder, 1991 and Laffitteina mengaudi (Astre, 1923); Fanrhapydionina flabelliformis Fleury, 2014 is rare. This zonation may be compared to the chronostratigraphic scale if one accepts the age assignments usually retained for larger benthic foraminifers of open sea, more ubiquitous, sometimes intercalated in the inner platform facies: • Orbitoides tissoti Schlumb., 1902 and Pseudosiderolites vidali (Douvillé, 1906) would be roughly of Campanian age, according to a large compilation due to Guˇsi´c and Jelaska (1990); • Orbitoides media (D’Arch., 1837) would appear approximately by mid-Campanian, according to the same authors; • Siderolites calcitrapoides LMK., 1801, Omphalocyclus macroporus LMK., 1816 and Orbitoides apiculata Schlumb., 1901 are usually considered as Maastrichtians. In these circumstances, O. tissoti and P. vidali observed prior to the first Rhapydioninidae levels (Mavrikas, 1993) would indicate the Campanian age of the CsB6a biozone. In addition, O. media observed in levels earlier or intercalated with CsB6b association (Luperto Sinni and Ricchetti, 1978; Chiocchini and

Mancinelli, 2001) indicates a high position in the Campanian. Finally, the biozone CsB7 is known superimposed to S. calcitrapoides, O. macroporus and O. apiculata (Fleury, 1973, 1980) which places it in the Maastrichtian. The Klokova massif, in Southern Continental Greece (Fig. 1, and Fleury, 1980 for more details), is the only known outcrop displaying a continuous succession of Upper Cretaceous carbonates of the Gavrovo-Tripolitza platform. Unfortunately, no significant fossils were found between the two Rhapydioninidae bearing levels, which are: - The older one including Pseudochubbina philippsoni, Cuvillierinella fluctuans nov. sp. and Cuvillierinella nov. sp. is supposed to indicate the CsB6a zone, mainly because of the absence of any CsB6b markers. The only positive fact would be the presence of P. philippsoni, congeneric with P. bruni, a companion of C. salentina in its type locality. - The younger one (irremediably covered in part) is a gathering of almost all members of the CsB6b zone, with Metacuvillierinella nov. gen. It precedes the levels with Rhapydionina gr. liburnica (recrystallized in this outcrop). Moreover, the Strontium Isotope Stratigraphy method brings some interesting new data (Vicedo et al., 2011). The age of the type locality of C. salentina would be 76.6 ± 0.6 MA, constraining to shift the CsB5–CsB6 boundary in an older position than in Fleury, 2014 (fig. 3). The type locality of C. pylosensis (in the vicinity of the type locality of Cyclopseudedomia smouti) would be about 71 MA, which is compatible with Fleury’s 2014 hypothesis. These data and assumptions are summarized in Fig. 2 that also gives a schematic picture of the changing conditions of sedimentation during the studied period (Landrein, 2001; Landrein et al., 2001); the essential feature is an important emersion widespread at the boundary between biozones CsB6 and CsB7. This event is interpreted as having caused the disappearance of the Rhapydionininae characterizing the CsB6 biozone. It could be assumed that the same type of event is at the origin of the discontinuity of settlements between the two divisions of the CsB6 zone; this assumption is new, justified by the fact that, previously, C. pylosensis and C. aff. pylosensis were not distinguished from C. salentina (see Fleury, 1974), but it must be emphasized that it is not based on actual field observations. Finally, it should be noted that, according to Cvetko et al., 2001 (fig. 2), in the island of Bra´c, Dalmatia, levels with Orbitoides media would precede an association with K. tergestina, M. apenninica and Reticulinella fleuryi. Such a piece of information is not compatible with those of Fig. 2, but the synthetic patterns of these authors show lateral passages of facies that could be a source of confusion; a confirmation is needed before this eventuality is considered as a fact. 5. Main features and nomenclature applied to the Rhapydioninidae As part of the superfamily Alveolinacea, the Rhapydioninidae share with the Alveolinidae a special mode of partitioning which characterizes the whole group: each chamber is divided

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Fig. 3. Architectural overview of a theoretical Cuvillierinellinae with examples from Cuvillierinella spp. A: Supposed test cut by two half-plans coincident with the coiling axis (axial section) and the equatorial plane (equatorial section). Note the succession of four stages: 1: streptospiral to milioline around the proloculus, eventually missing; 2: involute planispiral (chambers B1 and D); 3: pseudoevolute (chamber B2, partly involute, partly evolute); 4: evolute (chambers B3 and E-F, eventually cyclical). B: Isolated chambers from the various stages. B1: Chamber isolated from the involute stage (the cloisonnettes are here fundamentally perpendicular to the coiling axis). B2: Chamber isolated from the pseudoevolute stage (a transitory stage: the chamber being involved both in the involute and evolute stage). B3. Chamber isolated from the evolute stage (the cloisonnettes are here fundamentally parallel to the coiling axis). C: Actual centered equatorial section of C. salentina with cylindrical UUT (see Fig. 6.1) displaying the whole set of endoskeletal elements. Note the discrepancy between the orientation of the primary axis of openings (O1Ax) and primary and secondary chamberlets (Ch1Ax, Ch2Ax) in the last two chambers, as emphasized in reconstructions of figure E and F. D–F: Close-up on the endoskeleton organization of dissected chambers from various stages (D: involute planispiral, E: evolute cylindrical, F: evolute flabelliform), with informed actual sections (C–E: Cuvillierinella salentina, from Fig. 6.1, Fig. 6.7 and Fig. 5.9; F: C. aff. pylosensis, from Fig. 9.15). Abbreviations: Ce: Central endoskeleton; Ch1, Ch2: Primary and Secondary chamberlets; Cl: Cloisonnettes; O1, O2: Primary and Secondary openings; P: Proloculus; Pp: Preseptal pillars; Ps: Preseptal space; S: Septum.

into tubular chamberlets, parallel to direction of coiling, connected by an undivided volume located in the anterior part of the chamber, the preseptal passage, where pillars (preseptal pillars) appear in the advanced tests. The Rhapydioninidae are characterized by chambers presenting a special apparatus, a central thickening pierced by canals (the secondary chamberlets), originally parallel to the primary chamberlets, but capable of some complexity (the helicoidal structure of Fleury, 1979a); accessorily, it can be added that the tests, involute, subspherical or lenticular in early stage tend to uncoil as to form in both generations an evolute terminal part, cylindrical, flabelliform or eventually cyclical (Fig. 3). The nomenclature used below is mainly after Reichel (1936–37, 1964), Smout (1963), Fleury (2014 and previous works) with some alteration. Fig. 3 illustrates the main terms.

The theoretical shell (Fig. 3.A), streptospiral then planispiral, involute then evolute is made up of 3 sorts of chambers: crescent (or banana-) shaped (Fig. 3.B1 and D) in the involute stage, Y shaped (Fig. 3.B2) in the intermediate stage (pseudoevolute), part of a circle (Fig. 3.B3 and F) in the final evolute stage; a cylindrical terminal stage is equally extant in some species (Fig. 3.E); the general term for final pseudoevolute–evolute stages will be “Uncoiled Uniserial Termination”, abbreviated UUT afterwards. Every chamber is coated by a wall, of which the part in contact with the previous coil is classically called “basal layer”, not to be confused with the central endoskeleton. The wall becomes the septum (S) or apertural face, where it is covered by the next chamber. The pores that strew the septum are the openings or apertures; the main openings, arranged in one row at the edge of the septum, are called primary openings (O1) and,

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as a rule, secondary openings (O2) are scattered all over the central septum area; these secondary openings, like the whole set of chamberlets, are arranged in the direction of coiling, but the primary openings are oblique to this orientation and decentralized with regard to the corresponding chamberlets (Fig. 3.C, E, and F, Fig. 5.20, Fig. 7.2); this particular feature recalls in part the arrangement of the same elements in Rhapydionina (see Fleury, 2014), as witness of a certain kinship. The endoskeleton, inside the wall, is made up of cloisonnettes (Cl) (the term “septula”, frequently used, is too general to apply to such a special element, giving birth to the central endoskeleton), parallel to the direction of coiling, isolating primary (or cortical, or marginal) chamberlets (Ch1), born from the primary openings. The cloisonnettes fuse in the center of the chamber to form the central endoskeleton (Ce), pierced by more or less irregular secondary (or medullar, or central) chamberlets (Ch2), born from the secondary openings; they are sometimes arranged in layers separated by “floors”, sometimes disordered, the distance between them being often larger than their diameter, which is described by the expression “Scattered Secondary Chamberlets” (SSC). All these chamberlets are connected in the anterior part of the chamber by the preseptal space (Ps), crossed by eventual preseptal pillars (Pp) (“residual” pillars has no meaning, being the recollection of a long forsaken theory assuming that, issued from soritids, the rhapydioninids get their central endoskeleton from lateral fusing of fully mythical interseptal pillars). 6. Legitimacy of the genus Cuvillierinella Papetti and Tedeschi, 1965 This genus was one of the earliest Rhapydioninidae known from the Old World. The type species, C. salentina, described from Upper Cretaceous of Southern Italy, is the most widely represented but some other species will be described below. This genus is now accepted by recent authors (Vicedo et al., 2009, for example) but it can be useful to recall that, for several decades, European authors did not use it, to the benefit of Raadshoovenia Van Den Bold, 1946, a Cenozoic genus from Guatemala. This was not the result of a mistake, but it was due to lack of information. The type species of the American genus was only known through few explicit drawings, impossible to compare directly with the Italian species. The revelation of the likeness is due to De Castro (1971) who published several good sections from the type material of R. guatemalensis, the type species of the genus, obviously resembling C. salentina. Hamaoui and Fourcade (1973) soon supported that view, giving new sections of the type material of the same aspect. There was no more doubt about the precise analogy of the two taxa and, as the Cenozoic age of the American taxon seemed not assured, many authors admitted that homology was likely and, accordingly, relegated Cuvillierinella to the rank of junior synonym. Pêcheux was however to confirm (since 1984, in an unpublished work, known by some insiders) the Paleocene age of the American species, involving that the homology of the two taxa was not acquired and that phylogenetic systematics had to take it into account. Fleury and Fourcade (1990) took note of this

fact and recovered Cuvillierinella in its original rank; however, the question remained open for several authors, such as Vicedo et al., 2009, p. 234, who wrote: “It is unclear whether we are in presence of almost perfect homoeomorphs that substitute each other in time and on the two side of the Atlantic or if there is a unique, cosmopolitan group that survives the events at the Cretaceous-Paleogene boundary”. There are indeed a number of analogies between the two taxa. They reside in the following characters: – proloculus spherical, of about 0.1 mm in diameter; – first whorls quinqueloculine to streptospiral; – final whorls planispiral, eventually followed by a short UUT; – chamberlets arranged in one then several layers; – preseptal space traversed by pillars; – size of tests, in the order of 1 mm. It must be added that a third species from Upper Cretaceous of Central America could be described in the same terms (except the missing final UUT): “Borelis” cardenasensis Barker and Grimsdale, 1937, successively attributed to Chubbina Robinson, 1968, Praechubbina Fourcade and Fleury, 2001, Chubbinella Vicedo et al., 2013. Thus, we are in the presence of a structural theme, the meaning of which being open to interpretation. It could be a stable structural type, having managed a Cretaceous transatlantic migration, in one or the other direction, followed by a Paleocene persistence limited to the New World. Alternatively, this theme would be nothing but another example of the ontogenetic recapitulation of recurring intermediate link, long admitted and never doubted, from a group of Milioliacea, characterized by their quinqueloculine to streptospiral coiling and lack of endoskeleton, to the Alveolinacea, tending to the planispiral coiling (see Pêcheux, 2002) with endoskeleton. Examination of the context specific to each of these taxa seems to clarify this apparent dilemma (Fig. 4). • Praechubbina cardenasensis would be issued from Pseudonummoloculina pecheuxi Fourcade and Fleury, 2001, through the simple attendant species Praechubbina breviclaustra, P. streptospira and P. compressa, all described by Fourcade and Fleury, 2001 (see discrimination of these species in their fig. 6), and is likely to give birth to various species of the genus Chubbina; • Raadshoovenia guatemalensis is accompanied by “plain forms” (Pêcheux, 2002) and sometimes by Neomurciella butterlini Fleury and Fourcade, 1987, a credible descendant; • Cuvillierinella salentina is accompanied by “plain forms” (see below and Fig. 5) which indicate its local origin and completely planispiral forms going to diversify widely, at least within the genus Murciella Fourcade, 1966 in one part and the genus Cyclopseudedomia Fleury, 1974 in another one (see Fig. 13). Thus, each of the three taxa is included in a particular context, well localized in time and space, which leaves no place for doubt: the common Raadshoovenia–Cuvillierinella structural motif corresponds to ontogenetic recapitulation of phylogenesis, as suspected by Pêcheux (2002). A common milioline stock of origin is possible, comparable environmental conditions are probable, and it remains that a classification of

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“natural” inspiration cannot tie together in the same taxon these milestones of separate evolutionary lineages. Returning to Europe, we will examine the type species of Cuvillierinella from its locality of origin, then several populations of Greece and Spain which are more or less close to it, under the name of Cuvillierinella gr. salentina as a whole. Another genus (Metacuvillierinella nov. gen.), more monolithic, will be described from a new species, which presents the same basic characters but is distinguished by its morphology and its particular stratigraphic distribution, following a quite different evolutionary path. 7. Systematic paleontology Order Foraminifera Eichwald, 1830. Suborder Miliolina Delage and Hérouard, 1896. Superfamily Alveolinacea Ehrenberg, 1839. Family Rhapydioninidae Keijzer, 1945. Subfamily Cuvillierinellinae nov. subfamily. Type genus: Cuvillierinella Papetti and Tedeschi, 1965. Diagnosis: Euro-Asiatic Rhapydioninidae of Campanian– Maastrichtian age, streptospiraly coiled in their nepionic stages (or clearly issued from streptospiral ancestors) and planispiral or nearly in their terminal parts, distinct from all others by the subcylindrical volume of their primary chamberlets (sub circular shape of the transverse section), as shown by Fleury, 2014, fig. 19. All members are linked to Cuvillierinella of which the various morphotypes of the type species and other related species (C. perisalentina nov. sp. and C. fluctuans nov. sp.) lead to the distinct evolutionary tendencies followed by other genera: Murciella, Cyclopseudedomia, Metacuvillierinella nov. gen. and Pseudochubbina (Figs. 13 and 14). Remarks: This new subfamily results from the recognition of the clearly marked individuality of the structural model Rhapydionina–Fanrhapydionina (see Fleury, 2014), constituting the nominotypical subfamily Rhapydionininae Keijzer, 1945.

Fig. 4. Praechubbina cardenasensis (9–11) and Raadshoovenia guatemalensis (15–20) in the broad outlines of the American subfamilies Chubbininae (Campanian–Maastrichtian) and Neomurciellinae (Paleocene–Lower Eocene) (solely A tests). 1–2: Pseudonummoloculina pecheuxi (from the type locality; 2: holotype). 3–4: Praechubbina breviclaustra (from the type locality). 5–6: Praechubbina streptospira (from the type locality; 5: holotype). 7–8: Praechubbina compressa (from the type locality). 9–11: Praechubbina cardenasensis. 12: Chubbina macgillavryi. 13: Chubbina jamaicensis. 14: Chubbina fourcadei Vicedo et al., 2013. 15–20: Raadshoovenia guatemalensis; 15–17: “plain forms”, 18–20: “typical forms”. 21–22: Neomurciella butterlini (from the type locality). Scale bar: 1 mm. 1–9, 11: in Fourcade and Fleury, 2001. 10: interpretative drawing after Barker and Grimsdale, 1937, pl. 9, fig. 1. 12: new, from Mexico (EJF39–JJF1145). 13–14: new, from Mexico (EJF1–JJF1118, EJF64–JJF1158). 15, 19–20: new, from Mexico (EJF26–JJF1135). 16–17: interpretative drawings after Pêcheux, 2002. 18: interpretative drawings after Hamaoui and Fourcade, 1973. 21–22: in Fleury and Fourcade, 1987.

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The unity of the group comes from their likely common ancestor, which seems to be represented by the “plain forms” described from the type population of C. salentina (Fig. 5.11–16). In the same population, various morphotypes give evidence of the potentialities developed by other genera which lose the streptospiral initial coiling: –refinement of the endoskeletal mesh (“Murciella-like forms”) without or with very faint final uncoiling for Murciella; –shortening of the involute initial coil to the benefit of the final uncoiled terminal part associated to large proloculus and important dimorphism for Cyclopseudedomia. In another path, Metacuvillierinella, close to Cuvillierinella by its endoskeleton and its initial coiling, develops an advolute mode of coiling and has a very faint dimorphism. A little less resembling to C. salentina, the Pseudochubbina species seems to be linked to C. perisalentina nov. sp. through their persisting “slow axial rotation” and their “scattered secondary chamberlets”. Differences and resemblances: The Rhapydionininae, the nominotypical subfamily of partly the same period and same area are distinguished by their mainly planispiral tests (although the simplest or primitive forms are streptospiral in their nepionic stage), their important dimorphism of generations, and their particular shape of primary chamberlets, triangular or rectangular in transverse section (their radial dimension being always much larger than the distance between two adjacent cloisonnettes), with obliquely oriented primary openings at the very edge of the central endoskeleton (see Fleury, 2014, fig. 1 and 20). The Pseudedomiinae subfamily, of the same period, but rare in the Mediterranean area, is still ambiguous, because of the little known holotype of the type species; after the work of Smout (1963), it seems that the planispiral coiling throughout, the acute periphery of the A tests, the important dimorphism of generations and the very thin endoskeletal mesh (fine chamberlets, narrow preseptal passage, numerous preseptal pillars), giving birth to the helicoidal structure (Mavrikas et al., 1994) are the main characteristics of a group of species presently gathered in the genus Pseudedomia. An overview of the American subfamilies Chubbininae and Neomurciellinae is given in Fig. 4. Range: Probably Middle to Upper Campanian, eventually Lowermost Maastrichtian (zones CsB6a and b, see Fig. 2). Genus Cuvillierinella Papetti and Tedeschi, 1965. Type species: Cuvillierinella salentina Papetti and Tedeschi, 1965. Diagnosis: Euro-Asiatic Cuvillierinellinae; test with globular proloculus and flexostyle in A generation, milioline to exceptionally planispiral in the early stage, planispiral in the adult. Adult test involute, subspherical, slightly biombilicate, rounded in periphery in both generations. Final stage eventually unrolled, forming a short cylindrical evolute UUT in A generation, or a large pseudoevolute to evolute flabelliform (never cyclical or discoidal) terminal flange in B generation. Endoskeleton made up of more or less early tubular primary and secondary chamberlets, the diameter of the first ones being ordinary larger than the seconds. Preseptal passage relatively wide, with preseptal

pillars, rare in the young, disposed in one or several rows in the final chambers of B tests. Remarks: Despite the difficulties which were previously referred to, this genus is relatively well known, at least for its typical forms (the dominant type, described below). It brings together all the characters of the Rhapydioninidae family, including those of the endoskeleton and those of the coiling (from the streptospiral nepionic to the final uncoiled stage); tests are globose to weakly nautiloid, eventually flanked by a UUT, either cylindrical or flabelliform. All other genera in the family share this pattern and are distinguished by their evolutionary choices, on coiling (and uncoiling) mode, importance of the dimorphism of generations and detail of the endoskeletal organization. Murciella is the only genus of precise comparable morphology. It could make the matter of a long debate but, for the moment, a working definition of the difference between Cuvillierinella and Murciella would stress on the streptospiral young coiling and late endoskeleton with large mesh of Cuvillierinella, versus wholly planispiral coiling and early endoskeleton with fine mesh of Murciella. Range and occurrence: Middle to Upper Campanian and probably Lowermost Maastrichtian (Zones CsB6a and b, Fig. 2), in the innermost parts of carbonate platforms of Greece (including some eastern islands), Dalmatian islands of Croatia, Southern Italy, and Spain (Betic Cordilleras). Cuvillierinella salentina from its type locality. Several samples are available from the type locality (quarry at Vitigliano, Peninsula Salentina, Province of Lecce; see De Castro, 1990; see Fig. 1: spot I340). Part of them, collected by the author, are identified I120, I336, I342 and I343 (collection JJF186, 229, 232 and 233) from which only the first and last contain populations large enough to be exploited below; others, collected and lent to the author by P. De Castro, are identified by the PDC sign followed by De Castro sample number (kept in his own collection, in the University of Napoli). This material had been partially studied by De Castro (1988) showing C. salentina associated with tests of Murciella type. This important finding will be developed and discussed below. The available material comprises essentially A tests from which the following categories can be distinguished: – “Dominant type”, constituting the majority of the population, – “Plain forms”, less frequent, fairly close to miliolids, – “Murciella like” forms, very rare. B tests will be finally described; they are apparently insignificant as they are multiform, but it is conceivable that this is proper to a taxon yet close to its origins. “Dominant type” (Fig. 5.1–8). The dominant type constitutes the majority of the population; its analysis, faced with the holotype and some paratypes must offset the absence of previous general study of the type population. All of the measures mentioned below are recollected in Fig. 12; it provides, in particular for equatorial diameters, statistics on 50% of the measures (the largest), intended to

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Fig. 5. Cuvillierinella salentina from its type locality. 1–8: A tests of the “dominant type” (1: holotype). Note the streptospiral juvenile whorls (almost quinqueloculine in 1, 2 and 4), the flexostyle in 3 (although the first whorl is of miliolid type), the few partitions before the last involute whorl, the large endoskeletal mesh in the last whorl of axial sections (5–8) and preseptal pillars in the last chamber of 1. 9–10: Transverse sections of chambers from a cylindrical UUP (Uniserial Unrolled Part); note the preseptal pillars on 9 (see Fig. 3.E). 11–16: A tests of “plain forms”, characterized by their very few partitions; note the flexostyle on 11, 13 and probably 14. 17: An unusual A test with very tight coiling, note the flexostyle associated to a miliolid like first coil. 18–26: “Murciella like forms”. 18–21: A tests almost planispiral with large endoskeletal mesh (flexostyles on 19 and 21); in the last chambers of 20, note the oblique orientation of the primary openings, oblique to the direction of chamberlets. 22–26: A tests planispiral or nearly with early partitions and small endoskeletal mesh. 27–28: Transverse sections of chambers from cylindrical UUP, with small endoskeletal mesh (A or B tests). 29–31: B tests, relatively small, with large endoskeletal mesh, resembling the A tests of the “dominant type”, except their tiny compact milioline nepiont. 32: B test, very large, resembling the B tests of C. aff. pylosensis (Fig. 9.12, 14 and 22). 33–34: Large B tests with small endoskeletal mesh (see more B tests in Fig. 14). Scale bars = 1 mm. 1: Holotype, drawing after the original photograph in Papetti and Tedeschi, 1965. 2–4, 23 and 31: I343 (JJF233). 5–6, 8–22, 24–30 and 32–34: PDC7957, -7960, -7966 and -7874. 7: I336 (JJF229).

compensate for the presence of immature tests and the error related to axial sections measures that rarely correspond to the actual larger equatorial diameters. The proloculus is circular in section, of an internal diameter of 0.11 mm in the holotype. Our richer two populations correspond respectively to averages of 0.11 mm and 0.10 mm, from 0.05 up to 0.16; they are therefore compatible with the holotype. A flexostyle is ascertained in section Fig. 5.3. The first chambers of the holotype are streptospiral, quite close to the quinqueloculine, and devoid of endoskeleton, which is the case in most of our sections (Fig. 5.1–8). The following chambers are planispiral for about 2 turns in the holotype and our populations; at least, a part of the first whorl appears to be devoid of endoskeleton, but the second shows usually up to 2 levels of chamberlets. The cylindrical UUP of the holotype exists also in our populations, but is quite

rare (less than 10% of the sections) and thus can be considered as incidental, no more than the indication of an evolutionary tendency. The frame of the holotype and tests of our material is thick and the mesh of the endoskeleton is wide. An essential point lies in the shape of the primary chamberlets in section perpendicular to their axis, visible on some paratypes (Papetti and Tedeschi, fig. 5.1 and fig. 6.1–3) and here Fig. 5.5 and 9–10; the detail of their form and their size is variable, but they are always inscribable within a circle of which the dimension compared to the radius of the chamber is short; in other words, the layer of primary chamberlets fill a small place in the chamber, which is a remarkable difference with Rhapydionina, in particular (see Fleury, 2014). The preseptal space is relatively large. The preseptal pillars are rare and occur only very discreetly; they are clearly distinct in the last chamber of the holotype (Fig. 5.1) and

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in Fig. 5.9. Unrolled tests of some paratypes reach an equatorial diameter of 0.80–1.05 mm (Papetti and Tedeschi, fig. 6.2); the diameter average of the unrolled tests in the two main populations stands around 1 mm (maximum 1.12) and 0.90 (maximum 1.28). The diameter of the holotype (1.14 mm) is exceeded by the largest of our unrolled tests, reaching 1.88 mm. These differences are probably not significant, since the unrolled tests depicted Fig. 5.7–8 hit barely 1.2 mm, for example. Finally, note that for the richest two populations, the relationship between the size of the proloculus and the equatorial diameter is random. The material examined here contains so far a large number of sections that correspond to the type material. So, the species is well characterized and can be compared to other populations more or less resembling. But the type population includes deviant tests from the dominant type, which allow a better understanding of the evolutionary potentials of the species. “Plain forms” (Fig. 5.11–16). In the author’s own material and in three populations due to P. De Castro a number of tests have a comparable aspect to dominant forms but are distinguished by several characters: – a rather small size, – a streptospiral persistent coiling, – a late endoskeleton (in the last whorl and sometimes only in the last chamber), while the diameter of the proloculus of several of these tests is approaching the upper limit of the dominant type. A flexostyle is present (Fig. 5.11, 13 and probably 14). Therefore, it seems that they are no more than marginal specimens, which can evoke a close miliolid located at the origin of the species. “Murciella – like forms” (Fig. 5). According to the simple definition of the difference between Cuvillierinella and Murciella previously given and following De Castro (1988, pl. 7, fig. 2–9), three sorts of tests can be distinguished: • planispiral or nearly, with flexostyle, some tests show late partitions although endoskeletal mesh is wide, as in the dominant type (Fig. 5.17–21); • planispiral or nearly, some tests are equipped with early partitions and small endoskeletal mesh (Fig. 5.22–24); the transverse sections of Fig. 5.27–28 could correspond to such tests; • planispiral throughout and comprising small endoskeletal mesh, some rare and small tests looking like Murciella (Fig. 5.25–26). The B tests (Fig. 5.29–34) are not uncommon in our material, but as usual in little differentiated groups, difficult to characterize. They all have a juvenile part of quinqueloculine type, with about ten chambers or more, organized around a very small proloculus, followed by a streptospiral part whose diameter is close to the same part of the A tests (see also Vicedo et al., 2011, fig. 6.7–9). Some tests reach a diameter of about 1 mm, without unrolled part (Fig. 5.29–30); others of barely larger size have a cylindrical UUT, sometimes short (Fig. 5.31) sometimes

very important (Fig. 5.33–34). A test with large flabelliform final part (Fig. 5.32), exceptional, recalls C. aff. pylosensis (Fig. 9). The endoskeletal mesh recalls either Cuvillierinella type (Fig. 5.29–32) or Murciella type (Fig. 5.33–34). The implicit question raised by De Castro was the following: should the two associated extreme types of tests from these samples be considered as true representative of what is called Cuvillierinella and Murciella (Murciella becoming in this case a junior synonym of Cuvillierinella) or are we in the presence of a population showing the passage from one to the other taxon that should be distinguished? Fleury and Fourcade (1990, fig. 6A–B) had arbitrarily answered this question by favoring the second term of the alternative. We know now that the Murciella type is very exceptional within the type population of C. salentina and that, if the dominant type of this species is associated with Murciella cuvillieri in many populations, intermediate forms are always missing. In addition, C. salentina has his own younger successors in the CsB6b zone (C. pylosensis and C. aff. pylosensis) and M. cuvillieri may be observed sometimes alone, as for example in its type population (Fourcade, 1966, and author’s new observations on the type material), as well as in the one which is illustrated by Vicedo et al. (2009). As a result, we will accept that the “Murciella – like forms” are simply a testimony of a particular evolutionary trend, which will be developed by M. cuvillieri. We now confront this population to several other associations related to the same model. As they are not exactly identical to the type and each of them being unique, we call them Cuvillierinella gr. salentina as a whole. Cuvillierinella salentina from Greece associated to Murciella gr. cuvillieri. The XGP110 sample (together with XGP107 and 111, from the same locality) contains a population (Fig. 6.1–12) very similar to the dominant type of the Italian one. The proloculus of comparable size (0.11 mm), the sequence of the 3 coiling stages, the size of the tests and the massive frame recall the type population. However, the streptospiral stage is never typically quinqueloculine, and remarkably, the endoskeleton already appears as soon as this stage. The darker appearance of the whole test is probably related to a stronger development of the endoskeleton and a narrower preseptal space. There are no “plain forms” in the population; the concomitant Murciella gr. cuvillieri are well distinguished by their perfect planispiral coiling and thin endoskeleton mesh, although their proloculus has the same diameter. B tests display a flabelliform final stage (Fig. 6.11–12) with confused orientation of secondary chamberlets, testimony of a rough helicoidal structure (see Fleury, 1979a), of which it is so far the only testimony in the genus. The XGP153 population (Fig. 6.13–18) is partly similar to the preceding, considering the proloculus diameter (0.12 mm), the equatorial diameter (average 1.15 mm) and lack of typical quinqueloculine young stage; but the endoskeleton is never present in the central whorls and the UUT is very rare. The B tests are not known. The concomitant Murciella gr. cuvillieri are easily distinguished by their planispiral coiling associated

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Fig. 6. Cuvillierinella salentina from Greece and Spain. 1–12: Population XGP110. 1–9: tests of A generation, resembling the overall aspect of “dominant type” of the type population, almost quinqueloculine to streptospiral with relatively large endoskeletal mesh, but with remarkable early partitioning (particularly 3–4 and 7) and flexostyle in 8. 10: transverse section of chamber belonging to a cylindrical UUP. 11–12: nearly centered equatorial and axial sections of B tests; the oblique orientation of secondary chamberlets in 11 (especially in antepenultimate chamber) indicates a rough helicoidal structure. 13–18: Population XGP153. 13–17: tests of A generation resembling the preceding, but with later advent of partitioning. 18: transverse section of cylindrical UUP; note the various diameters of the secondary chamberlets. 19–31: Population EFG1. 19–20, 22–29: tests of A generation with nearly quinqueloculine early stage (note the flexostyle in 22 and 25) and large endoskeletal mesh. 21: transverse section of a cylindrical UUP, much alike section 18. 30–31: centered axial sections of B tests. 20 and 31 were previously illustrated by Hamaoui & Fourcade, 1973 (with erroneous magnification). Scale-bar: 1 mm.

to a fine endoskeleton mesh; there are no tests of “intermediate” morphology. The FPM249 population, from Eastern Mediterranean area (Astypalia Island, sample from courtesy of Phedon Marnelis and Michel Bonneau) provides another example of mixture. The rare C. salentina tests, often recrystallized in their central part are not illustrated; several proloculus are about 0.11 mm in diameter and the largest tests reach a diameter of 1–1.2 mm (the larger being a specimen with unrolled final part). No B tests have been observed. The associated M. gr. cuvillieri are easily distinguished by their planispiral coiling and fine endoskeleton mesh (see Fleury, 1979a, “M. aff. cuvillieri”, pl. 5.16–19 and 25–26). Cuvillierinella salentina from Spain (Fig. 6.19–31). This material was collected by Eric Fourcade, who gave a few photographs of Cuvillierinella salentina (“EF.G1” in

Hamaoui and Fourcade, 1973, pl. 2, fig. 1, 3 and 9; pl. 5, fig. 9–10) together with the accompanying Murciella gr. cuvillieri (in Hamaoui and Fourcade, pl. 4, fig. 3–9; pl. 5.2, 5). The locality is given as “Upper Senonian, Pantano de Camarillos (Province of Murcia), Betic Cordilleras (Spain)”. Eric Fourcade also gave the author the slices from a sample EF89-89 of the same provenance. They are named together as EFG1 (JJF1364-65). The material is interesting by its similarities and its differences from the Italian type. Like this one, the Spanish population has a juvenile stage streptospiral often close to the quinqueloculine, devoid of endoskeleton, then a planispiral stage with few secondary chamberlets, and a short UUT. The coiling appears relatively looser than in the Italian tests, but the main difference lies in the small size of the proloculus being 0.07 mm in diameter for the two populations. Plain forms do exist (Fig. 6.22 and 28), with flexostyle in 22. The B tests (Fig. 6.30–31) are differentiated through their quinqueloculine-like juvenile coiling; some of these tests would

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Fig. 7. Cuvillierinella perisalentina nov. sp. 1–12: centered (except 8) sections of A tests displaying the various modes of coiling. 2–3, 6–7, 9–10 and 12 appear planispiral or nearly throughout; section 1 (holotype) is axial around the proloculus and equatorial in adult stage; section 4 is axial around the proloculus, becomes equatorial, then axial in the last whorl; 5 is nearly equatorial in the young and axial in the adult; sections 4 and 11 present a twisted aspect revealing the irregular mode of coiling. Note the very early cloisonnettes in every axial section. Note the openings, clearly visible in two sections: on 2, the rectangle shows them off, the primary one being oblique to the orientation of chamberlets; on 10, downwards, the secondary ones interrupt the septum and are related to the chamberlets of next chamber. 8: section through a chamber of an UUT. The secondary chamberlets are scattered in a massive endoskeleton like in sections 4 and 10–12. The preseptal pillars are thick, tending to hide the preseptal space in equatorial sections (A and B tests); they are obvious in some axial sections (6, 10 and 17). 13–17: B tests, built around a very small proloculus and a tiny streptospiral young part (13b, 17); the involute adults resemble the A tests but are larger and followed by a large pseudoevolute terminal flange; the overall aspect is not far from B tests of C. salentina and C. aff. pylosensis, but the involute part is larger (see Fig. 14); 13b (enlarged central part of 13) and 17 (probably immature) show the miliolid-like nepionic stage. Scale bars = 1 mm, but 13b = 0, 25 mm. The figured sections are included in thin slices deposited in the paleontological collections of the University of Lille, Sciences and Technologies under the following numbers. Fig. 1: USTL 3057; Fig. 2, 7 and 9: USTL 3058; Fig. 3–5: USTL 3059 to USTL 3061; Fig. 8–17: USTL 3062 to USTL 70, successively.

not be different in size from the A tests, and others show a final flabelliform stage. M. gr. cuvillieri is here mixed in the population, but there is no ambiguity in the identification of the sections, both for the diameter of the proloculus in the average of 0.14 mm (18 measures) and for the size of the endoskeleton mesh, much finer in Murciella. Cuvillierinella perisalentina nov. sp. (Fig. 7). Diagnosis: Species of typical Cuvillierinella structure, with thick framework. A tests with rounded proloculus of an average diameter of 0.15 mm, with short flexostyle, involute biombilicate spiral test of 4–5 whorls, eventually followed by a short evolute part; coiling apparently planispiral but often showing a faint deviation of its axis (SAR), resulting in a streptospiral aspect of the whole test. B tests resembling the A, but larger, with miliolid nepionic stage and a final pseudoevolute (to evolute?) flange. Endoskeleton, very early, of mixed aspect, in part resembling somewhat Cuvillierinella with chamberlets separated by more

or less continuous “floors”, in part resembling Pseudochubbina, with secondary chamberlets more or less regularly scattered in a solid central endoskeleton (SSC apparatus). Description: • Test morphology, chambers arrangement. A tests: proloculus circular to oval in section, with an average diameter of 0.15 mm, followed by a flexostyle occupying about 1/3 to 1/2 of its circumference. The nepiont is streptospiral, never of quinqueloculine type. The first chambers around the proloculus form a planispiral-like coil that sometimes persists in the whole test, but sometimes show a deviation of the axis of rotation: sections of the axial type in the young whorls passing to an equatorial type, and vice versa; this is what is called Slow Axial Rotation. Consequently the tests tend to a nautiloid shape, biombilicated, with rounded periphery, eventually slightly twisted. They comprise 4–5 whorls, the height of which increases slowly, before the last chambers; the very last whorl, beyond 1 mm, participate in some rare cases in a

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short UUT, cylindrical (Fig. 7.1 and 9) or possibly flattened (Fig. 7.11); the absence of transverse sections in the examined material of this unrolled part indicates its rarity. Test diameter is between 0.40 and 1.92 mm (average 1.01); the 36 largest exceeding 1 mm (1.24 mm ± 0.32). B tests: around a small juvenile part apparently of milioline to streptospiral character, the tests are comparable to the A ones, of the same shape and apparently planispiral but somewhat larger, followed by an unrolled terminal part of pseudoevolute (to evolute?) type; their diameter is about 3, up to 5 mm; they are not extremely different from the B tests of C. salentina or even C. aff. pylosensis (see Fig. 14). • Endoskeleton. The partitions appear very early in both generations, immediately in contact with the proloculus A and very close to the young streptospiral part in B; in A tests, the second layer of chamberlets appears as early as the second whorl; increase in the number of secondary chamberlets appears about the fourth whorl. The two layers of chamberlets may appear in the involute stage isolated by a regular and continuous partition (“floor”) of the same curvature as the wall (Fig. 7.3, 7 and 11–12), but in the last whorl of several tests, while the height of the whorl has increased, and especially in the unrolled parts, the arrangement is obviously less regular (Fig. 7.4, 8 and 10–11) and evokes the ordering known in the genus Pseudochubbina, with chamberlets scattered in a massive endoskeleton (SSC). The openings are located, as it is the rule, in the alignment of the chamberlets; they are discernible in every equatorial section, especially in section Fig. 7.2 (the axis of the primary one oblique to the chamberlets orientation); the secondary ones are visible in axial section of Fig. 7.10. The preseptal pillars are not always very distinct, but are present in every equatorial section; due to their thickness, they tend to hide the preseptal space which is rather wide (about one fourth of the chamber length) in A tests (Fig. 7.1 and 9) as in B tests (Fig. 7.13–14). The overall aspect of both generations, the relatively large mesh of the endoskeleton make an attempt comparing the new species to Cuvillierinella salentina, C. pylosensis and C. aff. pylosensis species. In that case, the A tests of the new taxon, larger and more rounded, cannot be confused; besides, the SAR and the aspect of the SSC in the last whorls have no equivalent in these species. But, precisely, those last features induce a comparison to Pseudochubbina species. The simple visual comparison of Fig. 13 shows that confusion between the new species and the four Euro-Asiatic species is not possible: none of them presents an overall likeness to the new species. Thus, it is useless to start a detailed study at this level; but the association of these 2 characters existing, in the old world, only in these two genus invites to reflection. This will be done in the conclusions. Material: About 100 sections of various orientations, included in 30 slices from samples I340 and I341 (Coll. JJF230 and 231); 14 thin sections containing the illustrated sections are housed in the public paleontological collections of the University of Lille, 1. Sciences & Technology under the USTL n◦ 3057–3070.

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Holotype (Fig. 7.1): Centered equatorial section of an A test with a proloculus of 0.14 mm in diameter. The following involute spiral part comprises 4.5 whorls, appearing in the early ones cut by an axial plane and later by an equatorial plane: it follows that the coiling appears streptospiral in the whole. The last three chambers form a probably cylindrical UUT. The section is included in a slide on which are written the sample number (I341), the production number (4818), the author’s collection number (JJF231); its USTL number is 3057. Etymology: The prefix peri comes from the Greek and Latin; it is used in the meaning of in the surroundings of, close to, both corresponding to the geographic vicinity and morphologic aspect of the new species compared to the type species of the genus. Range and occurrence, type locality: The samples come from a quarry at Vitigliano (Southern Italy), localized and illustrated in detail by De Castro, 1990 (see also Fig. 1), from which the types of C. salentina and Pseudochubbina bruni De Castro, 1990 are also issued. The samples are not precisely from the quarry, but from slabs forming the walls of the surrounding fields. The age of this locality would be 76.6 ± 0.6 MA after Vicedo et al., 2011, but the species range is approximately Middle Campanian (CsB6a, Fig. 2). Cuvillierinella fluctuans nov. sp. (Fig. 8.1–9). Diagnosis: Tests of A generation with globular proloculus of 0.14 mm in average. Tests large for the genus, of about 3 whorls, the first being eventually streptospiral but alternatively planispiral as the rest of the shell, with eventual unrolled terminal part, either cylindrical or flabelliform. Endoskeleton early, appearing as soon as the first whorl; primary and secondary chamberlets forming an isodiametric network with polygonal mesh in the last whorl and the terminal part. No section of eventual B test has been observed. Description: - Test morphology. The proloculus of circular section of which the diameter is about 0.14 mm in the average is followed by a flexostyle of about half the circumference (Fig. 8.3). The first chambers are relatively large, either streptospiral (Fig. 8.2, 4–5 and 7), either planispiral (Fig. 8.3 and 6), some case being doubtful (Fig. 8.1); no relation seems to exist between the proloculus diameter and the mode of coiling. The whole tests comprise 2–3 whorls relatively loose and an unrolled terminal part of 2–3 chambers, either cylindrical (Fig. 8.5) or probably flabelliform (Fig. 8.6). The equatorial diameter is about 1.40 mm in the average, the largest of not unrolled specimens being 1.60 mm and unrolled specimens reaching 2 mm. The equatorial diameter seems a function of the proloculus diameter: it is about 1.10 mm for proloculus of about 0.10 versus 1.30 mm for proloculus of about 0.16. - Endoskeleton. The cloisonnettes appear as soon as the first whorl in several cases (Fig. 8.1–3 and 7); the central endoskeleton with secondary chamberlets exists in every case in the penultimate whorl and the last chambers exhibit a very particular endoskeleton made up of isodiametric primary and

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Fig. 8. Cuvillierinella fluctuans nov. sp. and Cuvillierinella sp. 1–7: Cuvillierinella fluctuans nov. sp. Tests of A generation; 3 and 6 are almost perfectly planispiral, 2, 4–5 and 7 (the holotype) show a distinct streptospiral early stage and 3–5 (6?) a flexostyle; the endoskeleton, very early in some case (1, 7), displays a network aspect, made up of somewhat polygonal isodiametric chamberlets (7–9). 10–13: Cuvillierinella sp.: A tests in centered sections, not unlike C. fluctuans nov. sp., but slightly streptospiral, with early partitioning and a tighter coiling. Scale bar = 1 mm. The figured sections are included in thin slices deposited in the paleontological collections of the University of Lille, Sciences and Technologies under the following numbers. Fig. 1–4: USTL 3071 to USTL 74; Fig. 5, 7 and 9: USTL 3075; Fig. 6 and 8: USTL 3076 and USTL 3077.

secondary chamberlets forming a unique network, nevertheless recalling some sections of C. salentina (Fig. 5.27–28). The preseptal space, of about 1/4 of the chamber length, can be seen in every equatorial section (especially in central whorls of Fig. 8.3); the preseptal pillars are well individualized in the two last chambers of Fig. 8.6. The openings are discernible in the chamber below the last one in Fig. 8.4. - Comparison. The A tests are the largest of all in the Cuvillierinella genus, even larger than C. perisalentina and C. aff. pylosensis, much younger; the proloculus, larger than in all populations of C. salentina is in the average of younger species. But the main distinctive feature is the network aspect of the endoskeleton with polygonal isodiametric mesh. Above this remarkable aspect, the special interest of the species is to show, in a different way from C. salentina, the transition from streptospiral to planispiral coiling, without any change in the main characters, both types of tests being identical, except their first whorl coiling. Material: 18 centered sections among about 40 sections of various orientations, included in 24 slices from the same sample (Coll. JJF-GKL420). 7 slices containing the illustrated sections are housed in the public paleontological collections of the University of Lille, 1. Sciences & Technology under the n◦ USTL 3071–3077. Holotype (Fig. 8.7): Test of A generation axially sectioned, showing the rounded proloculus (d = 0.14 mm), followed by about 3 whorls, the first one being streptospiral and the last two planispiral, for an equatorial diameter of 1.10 mm. The partitioning appears early in the streptospiral whorl and takes its network appearance with isodiametric polygonal mesh in the last cut chamber. The section comes from a slice identified by

three numbers GKL420 (sample), 7240 (fabric) and the author’s collection number JJF20. Its USTL number is 3075. Etymology: From the Latin fluctuans, in the meaning of fluctuating, varying, irresolute, as to make reference to the two coiling modes of A tests in the young. Range and occurrence: The type sample comes from the Klokova massif, South Continental Greece (see GKL locality in Fig. 1). The section is described by Fleury, 1980, p. 90–95. The age is approximately Middle Campanian (CsB6a zone). Cuvillierinella sp. (Fig. 8.10–13) This species, from sample GKL402 (Coll. JJF326), was presented by Fleury, 1979a (pl. 2, fig. 16–19), as “Murciella n. sp. 2”. In fact, the coiling of the few tests at disposition is almost planispiral, but section Fig. 8.11 shows a streptospiral nepiont. The relatively large diameter of the proloculus (about 0.14 mm), the early partitioning (Fig. 8.12–13) and the overall aspect of the tests indicate a probable affinity with C. fluctuans nov. sp., from which it is a neighbor in space and time (see Fig. 1, Klokova section). Cuvillierinella pylosensis Vicedo and Caus, 2011 2011 Cuvillierinella pylosensis Vicedo and Caus; Vicedo et al., p. 170, fig. 4.1–12, fig. 5.1–3 (not 4–5) This species was created from a sample very close to the type locality of Cyclopseudedomia smouti, in Greece, in the surroundings of Pylos–Methoni, Southern Peloponnesus (see Fig. 1, GGB locality; Fleury, 1974, p. 316). The authors found C.

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smouti and stated that their new species corresponded to “Cuvillierinella salentina” figured by Fleury (1974), associated to the type population of C. smouti. Nevertheless, the associations are different: their sample with C. smouti and C. pylosensis includes Rhapydionina fleuryi Vicedo and Caus, 2011, which does not exist in Fleury’s sample of 1974. According to Vicedo and Caus, the diameter of the A proloculus of C. pylosensis is 0.13–0.16 mm, and the equatorial diameter of the tests is 0.9–1.1 mm (involute tests), up to 1.7 mm (unrolled sections), although no complete equatorial section was illustrated. The involute stage is rather small; the first whorls are streptospiral but apparently never quinqueloculine; the last whorls are planispiral and end in a cylindrical UUT comprising up to 6 chambers. The partitioning is sometimes early and well developed in the adult tests, with a large mesh, the primary chamberlets being larger than the few secondary ones; the preseptal space is large, crossed by thick preseptal pillars. The authors presented some off-centered sections of probable B tests (fig. 5.1–3) interpreted after the sections of Fleury (1974); by the way, it must be pointed out that the known tests illustrated by Fleury (C. aff. pylosensis described below) are flabelliform with a thick central part and, as far as we know, the chambers never reach an annular stage. The distinction of C. pylosensis from C. salentina appears probably right, as the authors show a difference in the proloculus diameter and mainly a difference in age between the two species. But the arguments relating to the B tests are not suitable since the authors made an adventurous hypothesis on what is the B generation of their new species, and since the B tests of C. salentina and “C. salentina” of Fleury (1974) are not so different as they thought they are (compare Fig. 5.32 and Fig. 9.10–14 and 22). Moreover, the general aspect of the A tests being slightly different (see Fig. 13, for example), it seems more advisable to keep the distinction between the two morphotypes, and to wait for a better material when available. It remains that it will be always difficult to distinguish between poor representatives of C. salentina, C. pylosensis and C. aff. pylosensis in the absence of the associated species. The former material of Fleury is described below, under the provisional name of “C. aff. pylosensis”. Cuvillierinella aff. pylosensis Population GGB13 (Fig. 9.1–15), as described by Fleury (1974) under the name of “C. salentina”, is known only by a small number of sections (28 centered A tests and about 10 B tests) from a hundred of slices that were made to attempt to solve the then crucial problem of discrimination between large tests of C. smouti and the associated Cuvillierinella B tests. A tests display a proloculus of 0.13 mm on average, with a flexostyle (Fig. 9.6); the equatorial diameter is about 1 mm on average, up to 1.40 mm for involute tests, being hardly larger for unrolled tests. The young stage is streptospiral, very rarely almost quinqueloculine, probably devoid of partitioning; the planispiral chambers of the last whorl and the final unrolled chambers comprise secondary chamberlets (sometimes missing) of smaller

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diameter than the primary ones. B tests show a young part looking like the A tests, with quinqueloculine small nepionic chambers (Fig. 9.14); the adult part makes a pseudoevolute to evolute large flange of numerous chambers (up to 18), never cyclical. The cloisonnettes are continuous, without interruption, from one chamber to the next (Fig. 9.14). The secondary chamberlets are smaller than the primary ones, somewhat disordered (Fig. 9.11). The preseptal space is large, larger than in the associated C. smouti. The preseptal pillars are clearly visible in equatorial sections, forming one row (Fig. 3.F and 9.15) or two (Fig. 9.11). Population GGB511 (Fig. 9.16–23). This population is represented on 29 thin sections, also from the region of Pylos. No associated species allows to precisely define its age, but taking into account the fact that no observation of the CsB6a zone was made in this extensively explored region (see Fleury et al., 1978), the population is to be reasonably attributed to C. aff. pylosensis of the CsB6b zone. The proloculus diameter of A tests is 0.15 mm in the average, larger than in any population of C. salentina, the juvenile coiling is streptospiral, exceptionally planispiral, with a flexostyle (Fig. 9.17); the test diameter is barely larger than in the previous population (Fig. 12), and the UUT of some tests is probably flattened (Fig. 9.19). The partitioning appears early and the diameter of secondary chamberlets is smaller than the primary one. A large equatorial flabelliform section of a B test gives a particular interest to this population (Fig. 9.22); note that the tangential part of the section shows the continuous disposition of the cloisonnettes from one chamber to the next. Population GKL414 (Fig. 9.24–29) is known from 85 thin sections, comprising Murciella renzi, Rhapydionina dercourti (its type locality), Metacuvillierinella decastroi nov. gen., nov. sp. and Fleuryana adriatica of the CsB6b zone. The smaller tests being eventually confused with M. decastroi nov. gen., nov. sp., the diameter of the proloculus A is the only statistical data available; it is 0.12 mm on the average (9 measures), from 0.06 up to 0.14 mm; the juvenile part is either milioline or almost planispiral; the unrolled terminal part is cylindrical (Fig. 9.25–26) and probably slightly flattened too (Fig. 9.28). The diameter of secondary chamberlets appears smaller than the primary ones. No B test was observed. Population GGB345 (Fig. 9.30–33) is observed on 54 thin sections where M. renzi, C. smouti, M. decastroi nov. sp. and F. adriatica of the CsB6b zone are associated. As for GKL414, the possible confusion between the smaller tests of Cuvillierinella aff. pylosensis and M. decastroi nov. sp. prevents collection of statistical data, except the diameter of the A proloculus which is 0.16 mm on the average, from 0.12 up to 0.20 mm. The A tests are much resembling those of previous populations, with faint streptospiral juvenile part and eventual unrolled cylindrical terminal part. The secondary chamberlets diameter is smaller than the primary one. A section of a B test is known (Fig. 9.33), similar to section 10. The provisional name given to this species is certainly bound to very low knowledge of C. pylosensis, but also to its likeness with C. salentina. The one and only objective criterion would be the diameter of the proloculus, which seems relatively fragile (0.9–0.12 mm on average for the populations of

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Fig. 9. Cuvillierinella aff. pylosensis. 1–15: Population GGB13. 1–9: A tests, streptospiral to quinqueloculine in the young, with large endoskeletal mesh and short cylindrical UUP (2–3, 7–9). The species could hardly be distinguished from C. salentina without taking into account a statistic study of the proloculus diameter and seems different from C. pylosensis through its more important coiled part. 10–15: tests of B generation (after Fleury, 1974) partly recalling C. salentina (see Fig. 5.32); after a juvenile stage resembling the A tests (but a tiny compact clew), thicker than the flabelliform stage which is in no way made up of annular chambers (10, 12–14); the cloisonnettes appear continuous (without any interruption) from one chamber to the next on periphery of sections (see especially 14, to the left); secondary chamberlets appear scattered in a compact central endoskeleton (11); each section shows the distinctive large preseptal space; the section along the preseptal space of a chamber in 15 shows the elements labeled in Fig. 3.F. 16–23: Population GGB511. 16–21: A tests, streptospiral at first (with the exception of 17) then resembling in part the preceding population but with a probable short flabelliform uncoiled final stage (19). 22–23: B tests; note that there is no discontinuity of the cloisonnettes from one chamber to the next in periphery of 22 (especially to the top); 23: section perpendicular to the flabelliform flange, displaying large pillars more or less laterally fused, openings and cloisonnettes inside a single chamber, like 15. 24–29: Population GKL414 (associated with Metacuvillierinella decastroi nov. gen., nov. sp., see Fig. 11.11–16). A tests, displaying coiling modes from quinqueloculine to almost planispiral; the short UUT can be cylindrical (25–26) or perhaps compressed (28). 30–33: Population GGB345 (including Metacuvillierinella decastroi nov. gen., nov. sp., see Fig. 11.1–5). 30–32: A tests, typical, with cylindrical UUP; in 31 note the scattered secondary chamberlets of small diameter similar to those of 8–9. 33: B test much alike section 10. Scale bars = 1 mm.

C. salentina from Greece and Italy, 0.13–0.16 for C. pylosensis and 0.12–0.16 for the populations attributed above to C. aff. pylosensis). It remains that the biostratigraphic framework is the best justification for these distinctions and virtually the only way to try and distinguish these species from C. salentina in routine work.

Genus Metacuvillierinella nov. gen. Type species: Metacuvillierinella decastroi nov. sp. Diagnosis: Genus owning the characters of the family Rhapydioninidae in the sense of Fleury and Fourcade (1990), separated from all other genera by its compressed test in the equatorial

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plane and the apparently contradictory association of an advolute coiling to a total inability to final unrolling; the streptospiral coiling (frequently milioline in the young) is persistent, although decreasing in the adult, resulting in sigmoid axial section. The secondary chamberlets, often absent as a consequence of the flattened test, are of comparable size to primary ones. Both generations are represented by tests comparable in size and morphology, distinct only by their juvenile parts. The new genus is a member of the new subfamily Cuvillierinellinae through the typical shape of the primary chamberlets (circular in transverse section). Remarks: The type species of the new genus, M. decastroi nov. sp. resembles the Cuvillierinella species, such as the older C. salentina and the contemporary C. pylosensis and Cuvillierinella aff. pylosensis but differs mainly through a type of coiling unknown in the family until now, testimony of a special evolutionary pathway. This prevents from assuming M. decastroi to be a direct descendant of C. salentina, as we know it; but nothing prevents from supposing a common origin that would be rooted in the pre-Campanian time when, in the now well-known area of Western Europe, the ecologic conditions were not favorable to the spreading of the family. Metacuvillierinella (Figs. 10 and 11).

decastroi

nov.

gen.,

nov.

sp.

1967. Cuvillierinella salentina Papetti and Tedeschi; Horstmann, pl. 5.7 1977. Raadshoovenia guatemalensis Van Den Bold?; Fleury, pl. 1.1–11, 13–15 and 17–20 (not 12 and 16). 1986. Raadshoovenia sp.; Radoiˇci´c, pl. 3.2. 1990. Cuvillierinella sp. A; Fleury and Fourcade, fig. 6A.4, fig. 6B.4. Diagnosis: Species distinguished by a streptospiral to milioline nepionic coiling in any case, followed by a somewhat planispiral adult stage of the SAR type, advolute in both generations, resulting in flat tests, often sigmoid in axial section, never unrolled; B tests are hardly larger than the A ones and are only distinguished through their miliolid nepionic stage made up of small undivided chambers instead of the globular A proloculus. The central endoskeleton appears rather soon in the “planispiral” whorls; the endoskeletal mesh is large and the primary and secondary chamberlets are about the same diameter; in much flattened chambers, the central endoskeleton is reduced to a simple sagittal blade. Material: The type material consists of about 40 off-centered sections plus 10 centered sections of A tests and 4 B tests from sample GGB184 (Coll. JJF363). It comes from Skhiza Island, in the Pylos–Methoni area (see Fig. 1: GGB locality). The precise spot is given by Fleury, 1980 (locality “FB”, p. 112, fig. 36). 3 slices from this stock are housed in the paleontological collections of the University of Lille, 1. Sciences and Technology, under the n◦ USTL 3078–3080; several sections illustrated in Fig. 10 are embedded in slices which were previously deposited in Lyon University under the number FSL147804, -808, -810 and -814 (Fleury, 1979a). The figured material from complementary samples I250, GGB22, GGB345, GKL311, GKL314, GKL414

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and FTC66 consists of 26 slices numbered USTL 3081–3106 (details in Figs. 10–12). Holotype (Fig. 10.2): Centered axial section of an A test comprising a proloculus of 0.12 mm in diameter, several streptospiral chambers, and 3.5 advolute whorls of which the apparent planispiral coiling is denied by the sigmoid aspect of the whole section; the equatorial diameter is 1.88 mm and the axial diameter 0.28 mm. The central endoskeleton appears as soon as the first “planispiral” whorl, enclosing a few large primary chamberlets; the last chamber shows only one distinct secondary chamberlet, of about the same diameter as the primary ones. The section comes from a slide identified by three numbers GGB184 (sample), B2 (fabric) and JJF363 (collection); the USTL number is 3078. Etymology: In honor of Piero De Castro, a friendly colleague, eminent specialist of the Alveolinacea. Descriptions: Population GGB184 (type sample, Fig. 10.1–9). The centered sections of A tests constitute a very homogenous set. The proloculus, circular in section is 0.09 mm in diameter; the equatorial diameter of the tests is 1.70 mm on average. In axial sections, the thickness of the tests is hardly larger at periphery than along the axis, resulting in almost flat tests; the ratio between the axial diameter and the equatorial diameter being comprised between 1/6 and 1/10 (unspecified test, presumably B, Fig. 10.4), as a consequence of the advolute coiling. Some tests show a planispiral coiling, as a whole, but the sigmoid aspect of several sections reveals that the axis remains poorly stabilized as in the SAR mode. The A proloculus (with a probable long flexostyle in Fig. 10.9), is surrounded by several streptospiral chambers, milioline but never obviously quinqueloculine in this population, devoid of endoskeleton. The endoskeleton appears in the first “planispiral” whorl, and is ordinary still reduced to the cloisonnettes, convergent toward the center of the chamber, leaving little room to very few secondary chamberlets (Fig. 10.7), sometimes missing; the central endoskeleton is eventually reduced to a simple sagittal blade in the more compressed chambers (Fig. 10.8). The preseptal space takes up about one third or one fourth of the chamber length; no preseptal pillar is obvious in the sections at hand, they could be absent, as they are known to be generally dependent of a well-developed central endoskeleton. The B tests display a quinqueloculine juvenile part made up of undivided small chambers which fill about the same volume as the A proloculus (Fig. 10.3 and 6); the rest of the tests is about the same as the A tests in this population, but without any UUT. Population I250 (Fig. 10.10–18). This population, from Italy (see Fig. 1), is the most differentiated from the type population, but is still close to it. 33 thin sections allowed to make observations on 50 sections, of which 30 are centered. The A proloculus diameter is around 0.14 mm on average. The equatorial diameter is 1.90 mm on the average, up to 2.7 mm (Fig. 10.15); the ratio between the axial diameter is between 1/3 and 1/6 for the adult tests larger than 1.30 mm (13 measures); it must be kept in mind that such data are to be looked at in context: the axial diameter is dependent on the streptospiral coiling of first chambers and the equatorial diameter of axial sections is

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Fig. 10. Metacuvillierinella decastroi nov. gen, nov. sp. 1–9: Population GGB184. 1–2, 7–9: A tests (2: holotype) in centered sections, almost equatorial (1), and axial (2, 7–9) with flattened and more or less sigmoid tests; note the streptospiral coiling of early stages devoid of partitions and the rare secondary chamberlets of rather same diameter as the primary ones; sections 2 and 7–8 are perfect illustrations of advolute coiling. 3–6: tests of B generation (4–5 only probable), not very larger than the A tests, with quinqueloculine central part (3 and 6), isodiametric chamberlets and faint sigmoid shape of sections 4–6. 10–18: Population I250. 10–15: A tests in centered sections; note particularly the streptospiral nepionic coiling around the proloculus, the few secondary chamberlets of same diameter as the primary ones; 14–15: tests with an uncommon large diameter and a final tendency to unroll; a flexostyle is probable on 15. 16 and 18: probable B tests with remarkable mesh-like pattern of isodiametric chamberlets in the last chambers; 17: A test in centered axial section with no secondary chamberlets; 19–21: Population GGB22. 19: typical off-centered oblique section of an A test; 20–21: two ordinary axial sections of A tests. Scale bars = 1 mm. The figured sections are included in thin slices deposited in the paleontological collections of the University of Lille, Sciences and Technologies under the following numbers. Fig. 2: USTL 3078. Fig. 5 and 6: USTL 3079. Fig. 9–11: USTL 3080 to USTL 82. Fig. 13–21: USTL 3084 to USTL 3091. Fig. 1, 3, 4, 7 and 8 were deposited by Fleury (1976) in the collection of Lyon University under no. FSL147804, -808, -814, -810, respectively.

rarely the maximum diameter of the test. The other features fit well with the type population, except the following examples: Fig. 10.11 shows a tight coiling and a poorly developed endoskeleton; Fig. 10.12 displays a lately stabilized coiling; Fig. 10.14–15 shows two sections with equatorial diameters larger than in the type population; Fig. 10.15 reveals a probable flexostyle (salient protruding inside the proloculus, downward); Fig. 10.18 shows relatively well developed secondary chamberlets, of the same diameter as the primary ones. A single B test is probable (Fig. 10.16), poorly preserved but compatible with

the previous one. M. renzi, Fanrhapydionina aff. flabelliformis and Cuneolina sp. are associated with this population. Population GGB22 (Fig. 10.19–21). This population, from Pylos–Methoni area (locality “FA”, in Fleury, 1980, p. 112–114) has no remarkable features. Among the 25 sections observed on 20 slices, the proloculus diameter of the 8 centered A tests is 0.12 on the average, and the equatorial diameter lies in the average of the species. M. renzi and R. dercourti take part of the association. Population GGB345 (Fig. 11.1–5). From Pylos–Methoni area (see Fleury, 1977, p. 86). Despite the large number of available

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Fig. 11. Metacuvillierinella decastroi nov. gen., nov. sp. (associated with C. aff. pylosensis in every population). 1–5: Population GGB345 (see Fig. 9.30–33). 1–3, 5: typical tests of A generation in centered or nearly centered sections; note particularly the sigmoid aspect of 3, the few secondary chamberlets of about the same diameter than the primary ones; 4: remarkable B test, larger than the A tests but not unrolled, with tiny quinqueloculine nepionic stage; note the numerous secondary chamberlets of about the same diameter than the primary ones. 6–7: Population GKL311: A tests, an ordinary off-centered section (6) and a centered section of a large test (7). 8: Population GKL314: off-centered section of a probable B test, slightly flattened, with numerous secondary chamberlets. 9–10: Population FTC66: centered sections of A test; the smaller proloculus (9) is followed by a miliolid juvenile part, the larger (10) with long flexostyle is plainly streptospiral. 11–16: Population GKL414 (see Fig. 9.24–29): typical A tests with quinqueloculine to streptospiral initial coiling, flat in true axial section (11); secondary chamberlets are rare (and unusually small in 12) and preseptal pillars are only probable in antepenultimate and last chambers of 13. Scale bar = 1 mm. The figured sections are included in slices deposited in the paleontological collections of the University of Lille, Sciences and Technologies under the following numbers. Fig. 1 and 2: USTL 3092 and USTL 3093. Fig. 3 and 5: USTL 3094. Fig. 4–16: USTL 3095 to USTL 3106.

thin sections (54), the observed population is scarce, as most of the small sections cannot be definitively distinguished from C. aff. pylosensis (see Fig. 9.30–33). An excellent centered axial section of a B test (Fig. 11.4) summarizes all the information collected on the case, in particular on the homogeneity of the size of the different chamberlets, although the sigmoid aspect is lacking. M. renzi, C. smouti, Scandonea mediterranea De Castro, 1974 and F. adriatica are attendant on this population. Populations GKL311, GKL314 (Fig. 11.6–7 and 8, respectively). These populations, coming from a small outcrop on the eastern flank of the Klokova massif (see Fleury, 1977, p. 86), are rather poorly represented. They appear very similar to the type population. The centered sections of one of them (GKL311) give measures equivalent to the previous populations: in average, the proloculus diameter is 0.09 mm ± 0.02 (10 measures) and the equatorial diameter is 2 mm ± 0.20. A somewhat unusual test, large but without the sigmoid appearance, is shown in Fig. 11.7. The associated species are C. aff. pylosensis (probable), M. renzi, C. smouti, S. mediterranea and F. adriatica. Population GKL414 (Fig. 11.11–16). This is an abundant population, comprised in 85 slices from the Klokova massif (see Fleury, 1977, p. 88; 1980, fig. 29, p. 90, “F3”). The proloculus diameter of 34 centered sections is 0.10 mm ± 0.02, from 0.06

up to 0.14; the average of the equatorial diameters has not been established because of possible confusion with C. aff. pylosensis. The association comprises M. renzi, R. dercourti (type locality), S. mediterranea and F. adriatica. Population FTC66 (Fig. 11.9–10; JJF1044). This population comes from the Southern Peloponnese by courtesy of Franc¸ois Thiébault (localized in Fleury, 1977, p. 88); it is scarce but typical, with an average proloculus diameter of 0.12 mm ± 0.05 (5 measures), equatorial diameter of 1.51 mm ± 0.13 (9 measures) and the usual aspect of the species. M. renzi, R. gr. dercourti, C. hellenica, S. mediterranea and F. adriatica are identified in the population. Comparisons: Some characters, which make the extreme peculiarity of this species need not be considered here: no population of any different species may display good axial sections resembling the sections of the adult tests of the two generations. C. salentina may be difficult to distinguish from the immature test of the new species; an equatorial section of less than 1 mm is quite the same (Fig. 5.2–4, Fig. 6.2 and 4 versus central parts of Fig. 10.1, 11–12 and 14–15, Fig. 11.13–14); the same is true for oblique centered sections and, for example, sections Fig. 10.11–13 are given as belonging to the new species only because C. salentina is probably absent in the population.

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Fig. 12. Simplified comparative table of measurements of the studied populations. Every population is characterized by its sample identification (as in the text) and the number in the author collection (JJF). The graphics regarding Diameter of proloculus and Equatorial diameter summarize the statistical data given in the text. The equatorial diameter is given for 50% of the measures (the largest), as to avoid the immature individuals; unrolled tests are not taken into account; “doubtful measures” of GGB345 and GKL414 are due to possible confusion, in equatorial sections, between tests smaller than 1 mm of M. decastroi nov. sp. and Cuvillierinella aff. pylosensis. Abbreviations: M.: Murciella; Cycl.: Cyclopseudedomia; R.: Rhapydionina; Cuv.: Cuvillierinella; Pseudochub. spp. corresponds to Pseudochubbina bruni (I120 and I343) and P. philippsoni (GKL402, actually from GKL401, very near from 402, see Fig. 1).

C. pylosensis and C. aff. pylosensis may also present ambiguous small sections, but by lacking unrolled parts their tests are always smaller than those of the new species. Anyway, the B tests attest the difference. C. perisalentina nov. sp. and C. fluctuans nov. sp. share with the new species a difficulty to stabilize the planispiral coiling but are clearly distinguished through their involute coils and a whole different appearance. In short, although populations of the new species show some variability in the size of the proloculus, ranging between 0.09 and 0.14 mm on average (Fig. 12), well represented populations of other species cannot be confused with M. decastroi nov. gen., nov. sp. The equatorial diameter of the adult tests (devoid of UUT) is always larger than the tests of the known species of the genus Cuvillierinella, because of particular features which are the advolute coil, and compression in the sagittal plane resulting in flattened tests that also frequently take a sigmoid aspect. The partitioning happens relatively late; secondary chamberlets,

never numerous, are usually of a size comparable to primary ones. Tests of both generations share the same morphology and comparable size; even the B tests are never uncoiled and only characterized by their milioline nepiont made up of very small chambers. All of these characters form a whole which takes its meaning with regard to the distribution in time of this species compared to those which are the more closely resembling; as a result, the new species follows an original path that characterizes the new genus. The two references cited in the synonymy list should be noted again. The first one is, after Horstmann (1967), from Zanthe Island, an off-centered oblique unquestionable section, associated with some Murciella, among which M. gr. renzi is present; the sample comes from a reworked pebble included in a limestone of Eocene age. The second, after Radoiˇci´c (1986), is a 4 mm long (according to the given scale) off-centered section, which seems to be indubitable, assumed to be of a Campanian age, from a borehole in Iraq. The existence of this taxon in Iraq

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constitutes a new indication for exchanges of non-ubiquitous Larger Foraminifers between Western Asia and Mediterranean area by Campanian–Maastrichtian time that could not have been suspected a few decades ago. For example, Loftusia, and Pseudedomia, long considered as typical Eastern taxa are now known from Greece (Fleury et al., 1990; Mavrikas et al., 1994). Range and occurrence: In addition to the type material, several populations are observed from Greek and Italian various localities, (Fig. 1) numbered, I250, GKL311, -314, -414, GGB22, -345 and FTC66. The associated species are all from the CsB6b zone, approximately Upper Campanian (and partly Lowermost Maastrichtian?) in age. 8. Discussion and conclusion A classification tending to be natural implies a purely interpretative notion of the genus and must take into account the affinities, the evolutionary context more than the formal similarities, as the above mentioned questions about Raadshoovenia and Cuvillierinella recall us. In that view, there is no use to look for criteria that can be considered as basically of generic nature. A single character, constant in a population and absent from others, may characterize a species, but the notion of genus appeals to a group of related characters that indicate a new and original perspective, that is to say a particular evolutionary path within a group considered as well known. In general, these characters being independent from each other, operating at different speeds, the more the number of species reported in a genus, the more complex is the synthetic expression of the entity. This is the case of Cuvillierinella, of which the type species includes all types of characters that are barely the same in other species and are found in different combinations in the affiliated genera. This is what is developed below. 8.1. Genus Cuvillierinella This genus is the less specialized in the subfamily, a sort of protean model, which includes all potentialities implemented by the related genera. It is just a recollection of two sorts of characters already present in the older species (from CsB6a zone: C. salentina, C. perisalentina nov. sp. and C. fluctuans nov. sp.). On one hand stand the primitive characters, inherited from a miliolid source, such as small A proloculus, streptospiral coiling, few and/or late partitioning of large mesh, all of them still present in conservative younger species such as C. pylosensis and C. aff. pylosensis of the CsB6b zone. On the other hand are bound together the early beginnings of the evolutionary features that will characterize various evolutionary paths followed by other genera: –increase of A proloculus diameter (Murciella gr. renzi, Cyclopseudedomia), –loss of the streptospiral coiling (all genera, but SAR in Pseudochubbina and Metacuvillierinella nov. gen.), –thinning of the endoskeletal mesh with concomitant advent of the helicoidal structure (Murciella, Cyclopseudedomia), –acquisition of the SSC apparatus (Pseudochubbina), development of final unrolling (most of the genera, except Murciella gr. renzi), –extreme reduction of the

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spiral part to the benefit of the UUT (Cyclopseudedomia), and –increasing of the dimorphism of generation (Cyclopseudedomia, C. aff. pylosensis). Only one character is unknown from this panel: the advolute mode of coiling, which is the own property of Metacuvillierinella nov. gen. 8.2. Genus Metacuvillierinella nov. gen Consisting of a single species, this new genus is easy to define, being both partly similar to and partly different from Cuvillierinella. It looks like Cuvillierinella through its streptospiral juvenile stage, its wide endoskeleton mesh and its large preseptal space, so that the equatorial sections may be indistinguishable. It differs by its advolute type of coiling combined with the SAR, absolute resistance to the terminal unrolling and very low dimorphism of generations. It seems that if its origin may not be very different from that of Cuvillierinella, its future is unlikely, as most of the populations of the CsB6b zone, disappearing before the next zone. It is possible that the range of proloculus size of the various populations (Fig. 12) indicates an evolutionary trend but, for the present time, their relative age within the CsB6b zone cannot be determined. 8.3. An overview of the Cuvillierinella salentina group and its relatives Figs. 13 and 14 present the synthesis of observations and assumptions previously mentioned. The Cuvillierinella genus lies in a central position in both figures, because of multiple potentialities expressed by the many variants of C. salentina. They open many assumptions of filiations and in particular cousinhood, since this is probably to its milioline ancestors that we should be able to go back, in order to capture the precise relationships between the various species. The only track that comes to the mind would be that of a Miliolid group including both a streptospiral (preferably quinqueloculine) coiling and a flexostyle, an association which is shown by many sections of this work (Fig. 5.3 and 17, for example). Does such a group exist? 8.3.1. Taxa of the CsB6a zone On the one hand, if C. fluctuans, still little known, seems unlikely to pave a way for the future, C. perisalentina is more promising. Indeed, by its involute coiling of which the axis remains poorly stabilized (SAR) and its scattered secondary chamberlets (SSC) it strongly evokes the species of the genus Pseudochubbina. We know that A tests of P. kassabi and P. philippsoni have unrolled terminal parts, but it does not seem to be the case of P. bruni and P. globularis (according to De Castro, 1990), but B tests are probably not unrolled: they are still unknown in P. bruni and P. kassabi and the off-centered sections doubtfully attributed to this generation in P. globularis and P. philippsoni by their authors would correspond to subglobose tests. It is therefore very difficult to admit a direct relationship of these two taxa and it is more likely that they simply share a common origin. It should be noted that, although uncertain, this hypothesis of relationship of Pseudochubbina with any other taxon is the first to be supported.

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Fig. 13. A visual synopsis of the Cuvillierinellinae nov. sub family (A tests), with assumed relations between the various species in their stratigraphic framework. The species and genera are distributed according to their age (CsB6a and b). The links drawn between some of them do not imply any direct succession but indicate existence of some characters which are supposed to be the testimony of common origin (for example, M. decastroi nov. sp. is not supposed to be a descendant of C. salentina but could be considered as a cousin). All the species of the same biozone are supposed to be of same age, as far as we presently know (for example, the 4 species of Pseudochubbina are supposed of the same age, whatever their relative position, due to drawing constraint). The CsB6a–CsB6b limit is supposed to be related to a subaerial exposure (like the CsB6–CsB7 limit, see Fig. 2), because of the drastic change of population, although there is no actual proof of it. Scale bar = 1 mm. Cuvillierinella: C. salentina from Fig. 5, plus 2 sections from I343 sample; C. perisalentina from Fig. 7; C. fluctuans from Fig. 8; C. aff. pylosensis from Fig. 9; C. pylosensis after Vicedo et al., 2011. Pseudochubbina: P. bruni from type locality (I121–JJF187); P. globularis (Smout, 1963) from this author, type sample; P. kassabi De Castro, 1990, from this author, type sample; P. philippsoni after Fleury, 1977. Murciella: M. cuvillieri after Fourcade, 1966; M. renzi and M. ovoidea both after Fleury, 1979a. Cyclopseudedomia: C. hellenica after Fleury, 1979b; C. smouti after Fleury, 1974; C. klokovaensis after Fleury, 1979a. M. decastroi nov. gen. nov. sp. from Fig. 10.

On the other hand, regarding Murciella cuvillieri, its proximity in space and time with the populations of C. salentina, together with the existence of variants of the “Murciella – type” in the type population of C. salentina, lead to admit a very close relationship between the two taxa; however, this must be further examined in depth by the study of populations of M. gr. cuvillieri, whose B tests are still unknown. It will be recalled that if the two species are often associated with previously described populations, they are in every case clearly distinct and the type population of M. cuvillieri (as well as the one figured by Vicedo et al., 2009), includes no Cuvillierinella.

8.3.2. Taxa of the CsB6b zone Murciella gr. renzi raises quite different questions than M. gr. cuvillieri. The helicoidal structure, very well expressed in this group of species and recognized at rough state in M. cuvillieri (Fleury, 1979a; Fleury and Fourcade, 1987, pl. 3.11) is at the origin of the generic identification, but we know now that it is an evolutionary feature, appearing in the most advanced species of several independent lines (Fleury and Fourcade, 1990). As M. gr. renzi species (M. renzi and M. ovoidea in Fig. 13) are refractory to the final unrolling it is not certain that a direct filiation exists between these two groups of species; this is why they are separately linked to C. salentina in Fig. 13. This

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Fig. 14. Comprehensive view of the known B tests of Cuvillierinella and Metacuvillierinella species. The difference between the two genera is striking: at same size, the Cuvillierinella unrolled stage, cylindrical or flabelliform in C. salentina and always flabelliform in C. aff. pylosensis, has no equivalent in Metacuvillierinella. Note the large preseptal space of the Cuvillierinella tests, particularly obvious in these sections. Scale bar = 1 mm. C. salentina, after Fig. 5 and 6, plus several sections from the same populations. C. perisalentina nov. sp., from Fig. 7. C. aff. pylosensis, from Fig. 9, plus several sections from GGB13 (Fleury, 1974). M. decastroi nov. gen., nov. sp., from Fig. 10 and 11.

question should be considered in a setting both broader and more accurate than before, but it does not seem doubtful that the origin of these species must be situated in the vicinity of C. salentina. About the species of the genus Cyclopseudedomia, there are few questions since the description of C. hellenica by Fleury, 1979b. This species is resolutely planispiral but still close to Cuvillierinella by its relatively small proloculus (compared to other species of the genus), its wide preseptal space and its moderate final unrolling. It paves the way which is followed by the other species of the genus, more specialized and developing a more or less completed helicoidal structure, but its origin appears not doubtful. Cuvillierinella aff. pylosensis, little different from C. salentina, but younger, with larger proloculus, is obviously a direct descendant of this species. This is probably the same for C. pylosensis, although it remains too little known for comments. Finally, the posterity, or at least the kinship, of Cuvillierinella salentina proves to be richer and more abundant that one could imagine a few years ago. This is mainly due to the action of Piero De Castro who dared to deepen a matter apparently quite simple. Thus, it becomes clear that too simple concepts cannot account for the reality of the living; it may be inaccessible for long to us, but the lesson to draw is that only persevering work can help us approaching it.

Disclosure of interest The author declares that he has no competing interest. Acknowledgments The author is much indebted to Piero De Castro, who asked good questions about Cuvillierinella. In particular the author owes to him the loan of an important material from the type locality of C. salentina and expresses his deep appreciation and gratitude. A. Zambetakis and V. Vicedo provided constructive remarks on the initial text of the manuscript. References Barker, R.W., Grimsdale, T.F., 1937. Studies of Mexican Fossil Foraminifera. Annals and Magazine of Natural History London 10 (19), 110, 161–178. Chiocchini, M., Mancinelli, A., 2001. Sivasella monolateralis Sirel & Gundunz, 1978 (Foraminiferida) in the Maastrichtian of Latium (Italy). Revue de Micropaléontologie 44 (4), 267–277. Cvetko, B., Guˇsi´c, I., Jelaska, V., Buckovi´c, D., 2001. Stratigraphy and microfacies of the Upper Cretaceous Pucisca Formation, Island of Braˇc. Cretaceous Research 22, 591–613. De Castro, P., 1971. Ossevazioni su Raadshoovenia Van Den Bold e i suoi rapporti col nuovo genere Scandonea (Foraminiferida, Miliolacea). Bolletino della Società Naturalisti Napoli 80, 161–236. De Castro, P., 1988. Les Alvéolines du Crétacé d’Italie. Revue de Paléobiologie vol. sp. 2, Benthos’86, 401–416.

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