Citation: Serra-Kiel, J., Vicedo, V., Baceta, J.I., Bernaola, G., Robador, A., 2020. Paleocene Larger Foraminifera from the Pyrenean Basin with a recalibration of the Paleocene Shallow Benthic Zones. Geologica Acta, 18.8 , 1-69, I-III.
DOI: 10.1344/GeologicaActa2020.18.8

Paleocene Larger Foraminifera from the Pyrenean Basin with a
recalibration of the Paleocene Shallow Benthic Zones

J. Serra-Kiel1†

V. Vicedo2*

J.I. Baceta3

G. Bernaola4

A. Robador5

Universitat de Barcelona, Facultat de Ciències de la Terra
Departament de Dinàmica de la Terra i de l’Oceà, Carrer de Martí Franquès s/n, 08028
Barcelona, Spain.
1

Museu de Ciències Naturals de Barcelona, Departament de Paleontologia
Passeig Picasso s/n, 08003 Barcelona, Spain. E-mail: vvicedov@bcn.cat
2

3

Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU)
48080 Bilbao, Spain. E-mail: juanignacio.baceta@ehu.eus

Deparment of Mining-Metallurgy Engineering and Material Science, Faculty of Engineering in Bilbao, University of the Basque
Country (UPV/EHU)
Plaza Ingeniero Torres Quevedo 1, Bilbao, Spain. E-mail: gilen.bernaola@ehu.eus

4

Instituto Geológico y Minero de España
Ríos Rosas 23, 28003 Madrid, Spain. E-mail: a.robador@igme.es
5

* Corresponding author

ABSTRACT

A taxonomic study of the Paleocene larger foraminifera from the Pyrenean basin has led to the description of
sixty taxa including two new species: Alveolina korresensis and Valvulineria bacetai. In this work, we present
a chronostratigraphic recalibration of the Paleocene Shallow Benthic Zones SBZ 1 to SBZ 4 based on correlation
with calcareous nannofossil and planktic foraminifera biozones, all integrated within the stratigraphic framework
of Paleocene platform to basin depositional sequences established for the whole Pyrenean domain. The samples
were collected in autochtonous and parautochtonous deposits from ten key stratigraphic sections, representative of
coastal to platform margin depositional settings. The results from two sections representing base of slope facies
with intercalations of calcareous turbidites, which include penecontemporaneous platform-derived biota have been
integrated in the study. The regional chronostratigraphic framework is derived from magneto-biochronological
studies carried out in the Zumaia section, the global reference section for the Danian-Selandian and SelandianThanetian GSSPs. A new calibration of the Paleocene SBZs is proposed. The SBZ 1 is constrained to the first
1.09m.y. of the Paleocene; this first Paleogene biozone lacks distinct larger foraminiferal markers and thus is
defined by an association of non-exclusive taxa composed of Valvulineria patalaensis, Stomatorbina? binkhorsti,
Planorbulina? antiqua and Bangiana hanseni. The SBZ 2 now appears as the biozone encompassing most of the
Danian stage (from ca. 64.9m.a. to 61.6m.a.), and is characterized by the association of Haymanella elongata,
Haymanella paleocenica, Kayseriella decastroi, Rotospirella conica, Pyrenerotalia depressa, Elazigina
 J. Serra-Kiel, V. Vicedo, J.I. Baceta, G. Bernaola, A. Robador, 2020 CC BY-SA
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Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

dienii, Ornatononion moorkensii and Paralockhartia eos. The SBZ 2-SBZ 3 boundary coincides with the base
of the Selandian stage (ca. 61.6m.a.). The SBZ 3 biozone is defined by the occurrence of Glomalveolina primaeva,
Periloculina slovenica, Vania anatolica, Coskinon rajkae, Fallotella alavensis, Cribrobulimina carniolica,
Miscellanea yvettae, Miscellanea juliettae, Miscellanites primitivus, Miscellanites minutus, Ranikothalia
soldadensis, “Operculina” heberti and Discocyclina seunesi. The SBZ 3-SBZ 4 boundary is now ascribed to ca.
57.2m.a. The SBZ 4 biozone appears characterized by Glomalveolina levis, Alveolina korresensis, Hottingerina
lukasi, Daviesina garumnensis, Assilina yvettae, Assilina azilensis and Nummulites catari. The SBZ 4-SBZ 5
boundary is placed at the Paleocene-Eocene boundary event (ca. 56.0m.a.).
KEYWORDS

Larger foraminifera. Biostratigraphy. SBZ. Pyrenean basin. Paleocene.

INTRODUCTION
The earliest systematic studies on larger foraminifera
from the Paleocene of the Pyrenees are those by Mangin
(1954, 1955, 1960, 1966), Hottinger (1960), Caus et al.
(1980), Leppig (1988a, b) and Schaub (1981) in the SouthPyrenean domain and by Villatte (1962, 1969), Tambareau
(1966, 1972) and Tambareau and Villatte (1968, 1974) in
the North-Pyrenean realm.
The study of Paleocene larger foraminifera from the
Pyrenean basin received a major boost with the development
of the International Geological Correlation Programme
(IGCP) Project 286 “Early Paleogene Benthos” This project
.
organized two workshops in the Pyrenean basin. The first
was hold in Jaca (Spain), in October 1990. At this meeting,
several papers concerning stratigraphy, biostratigraphy and
sequential analysis of Paleocene outcrops in the Southern
Pyrenees were presented by Robador (1991), Serra-Kiel
et al. (1991), Robador et al. (1991a, b). The second
workshop was organized in Aspet (France), September
1994, with contributions concerning the stratigraphy and
biostratigraphy of Paleocene successions by Tambareau et
al. (1994a, b, c).

environments to platform-slope and hemipelagic
environments. In both theses, biostratigraphy was achieved
through larger foraminifera in the carbonate platform
environments, and planktonic foraminifera and calcareous
nannoplankton in outer platform and hemipelagic
environments. These correlations are reported in many
studies such as Baceta et al. (2004, 2005, 2006a, b, 2011),
Bernaola (2002, 2007), Bernaola and Nuño-Arana (2006),
Bernaola et al. (2007, 2009), Orue-Etxebarria et al. (1996,
2001), Pujalte et al. (1994, 1998a, b, 2000a, b, 2002, 2009a,
b, 2011, 2014, 2016) and Serra-Kiel et al. (1994).
These stratigraphic contributions were used by Baceta et
al. (2004) to define the depositional sequences (DS) called
Ma-Da, Da-1, Da-2, Se/Th-1 and Th-2, which encompass
the entire Paleocene, and were later incorporated into
various publications including a synthesis paper leading to
the formal definition of the Global Stratotype Sections and
Points for the bases of the Selandian and Thanetian stages
(Schmitz et al. 2011).

The first zonation of larger foraminifera of the
Paleocene series was developed by Hottinger (1960) based
on alveolinids. This biozonation included the G. primaeva
Zone for the lower Thanetian and the G. levis Zone for
the upper Thanetian. Later, Schaub (1981) characterized
the Assilina yvettae Zone for the late Thanetian based on
nummulitids. As one of the main conclusions of the IGCP
Project 286 ”Early Paleogene Benthos” the first integrated
,
biostratigraphic zonation for Paleogene platform series, the
Paleogene Shallow Benthic Zones (SBZ), was published by
Serra-Kiel et al. (1998).

The position of the Paleocene/Eocene boundary
in shallow platform successions at the base of the
Ilerdian stage has been of wide use in studies on larger
foraminifera from the whole Tethyan domain over the last
five decades. Several recent studies have demonstrated
that the Carbon Isotopic Event (CIE) that globally marks
the Paleocene-Eocene boundary accurately matches the
base of the Ilerdian in the stratotype and parastratotype
sections (Pujalte et al., 2003, 2009a, b; Schmitz and
Pujalte, 2003)This points out that the Ilerdian is indeed an
appropriate chronostratigraphic stage, at least at the scale
of the Tethyan realm, useful to characterize the shallow
platform successions deposited during the early Ypresian
stage, as defined by the International Commission on
Stratigraphy.

The PhD theses of Baceta (1996) and Robador (2005)
contributed to the stratigraphic and sedimentologic
knowledge of the Paleocene of the South Pyrenean realm.
Both authors studied and correlated numerous stratigraphic
sections from the continental-marine transitional

In this context, we present a new systematic study of
Paleocene larger foraminifera with the description of two
new species and a recalibration of the stratigraphic extent of
several taxa. With this updated database, we propose a new
recalibration of the SBZs from SBZ 1 to SBZ 4.

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MATERIAL AND METHODS
The study is based on the biostratigraphic analysis
of twelve sections of the Paleocene studied across the
southern Pyrenean domain, between Aragón and the
Basque Country regions (Fig. 1). Ten sections (Serraduy,
Campo, Tena, Garralda, Urrobi, Andia, Lizarraga,
Urbassa Pass, Leortza and Korres) are representative of
different settings across the shallow platform domain and,
consequently, are mostly made of shallow-water carbonate
deposits. The two remaining sections, Aixola-Ermua and
Monte Urko, represent base of slope areas and contain
hemipelagic deposits with intercalations of shallowderived resedimented deposits. All sections were logged
and sampled during different surveys carried out between
1990 and 2010. For the analysis of shallow benthic larger
foraminifera, a numer between fifty and hundred selected
hand-sized rock pieces were collected from each section.
Their study in the lab involved slabbing and selection of
specific areas for thin section preparation. More than one

Paleocene LBF from the Pyrenean Basin

thousand and five hundred individual thin sections of
standard size were processed and analyzed from all sections,
using petrographic microscope and binocular lens with
attached digital camera from the University of Barcelona
and the University of the Basque Country. Microscopic
analyses involved detailed identification and description
of the different taxa of benthic foraminifera, either
preserved in axial, equatorial and oblique sections, largely
based on digital images taken at different magnifications.
With the aim of simplification, only those samples and
corresponding thin sections with relevant content in larger
foraminifera were considered from each individual section
(see below). The thin sections studied and the associated
series of digital images are housed in the collections of the
Instituto Geológico y Minero de España, the Department
of Stratigraphy and Paleontology, University of the Basque
Country (UPV/EHU), the Department of Earth and
Ocean Dynamics, University of Barcelona (UB) and the
Department of Palaeontology, Museu de Ciències Naturals
de Barcelona (see Appendix I). The Josep Serra collection

FIGURE 1. Geological setting of the study area. A) Location of the Pyrenean basin in a general paleogeographic map of Western Europe. B) Paleocene
paleogeography and outcrop map of the Pyrenean basin. C) Location of the twelve reference sections for shallow benthic foraminifers studied.

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J. Serra-Kiel et al.

is now provisionally hosted in the Department of Earth and
Ocean Dynamics, University of Barcelona (UB) but will
possibly be transferred to the Museu de Ciències Naturals
de Barcelona.
The study also involved detailed integration of all
previous biostratigraphic data from calcareous nannofossil
and planktic foraminifera from the sections of Lizarraga,
Andia, Campo, Aixola-Ermua and Zumaia (Baceta, 1996;
Baceta et al., 2005; Bernaola, 2002, 2007; Bernaola et al.,
2006, 2007, 2009; Orue-Etxebarria et al., 2001; Pujalte et
al., 1993, 1994) and new data of calcareous nannofossils
from the Danian and Selandian of Aixola-Ermua and the
Danian of Monte Urko sections. These new data involved
the analysis of forty-six samples, seventeen from AixolaErmua section and twenty-nine from Monte Urko section.
The standard Paleogene scheme of Martini (1971), with
the amendments introduced by Perch-Nielsen (1981,
1985) regarding NP1/NP2 and NP2/NP3 zonal boundaries
and the more recent biostratigraphic scheme proposed by
Agnini et al. (2014) have been used for zonal identification.
The combination of these two biozone schemes provides
seventeen calcareous nannofossil zone boundary markers
for the Paleocene and thus accurate age constraints for the
stratigraphic succession to be established.
Geological and stratigraphic settings
In the Pyrenees, the Paleocene was a time of subdued
tectonism, which developed under a semiarid to arid climate,
globally rising sea levels and reduced siliciclastic supplies
from the neighbouring emerged lands (Baceta, 1996;
Baceta et al., 2004; Pujalte et al., 1993). The combined
effect of these general conditions led to a transgression
recorded across the whole Pyrenean embayment (Baceta,
1996; Baceta et al., 2004, 2006a, b, 2011; Pujalte et al.,
2000a, b; Robador, 2008). The general paleogeography
during this time consisted of a large sub-tropical marine
embayment opening to the paleo-Bay of Biscay and the
North Atlantic, with a 1000-1500m deep central basin
surrounded on its north, south and east sides by shallow
marine areas, in turn, flanked by broad continental alluvial
plains (Baceta, 1996; Fig. 1A, B). The shallow marine areas
were largely characterized by the deposition of skeletal
and non-skeletal carbonate sediments, forming platform
systems that in most sectors reached 30 to 40km across dip
and reached as much as 300-400m in composite thickness.
These carbonate platforms originally covered a total area of
at least 40,000km2.
The Paleocene platform successions from the Pyrenees
consist mainly of shallow-water limestones deposited
on beaches, tidal flats, lagoons, shoals and a variety of
reef constructions, but also include minor proportions of
siliciclastics and evaporites (Baceta et al., 2004, 2011).

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Paleocene LBF from the Pyrenean Basin

Pene-contemporaneous dolomitization likely produced by
evaporative processes was particularly important during the
Danian and middle Thanetian, and karstification took place
during the Selandian, which largely represents a period
of relative sea level lowstand (Baceta et al., 2001, 2007),
and during the Paleocene-Eocene boundary (Pujalte et al.,
2011, 2014). Basinwards, shallow carbonate platforms pass
sharply, through narrow slopes, to hemipelagic deposits, in
the form of rhythmic alternations of limestone, marlstone
and marl, locally with intercalations of thin calcareous,
siliciclastic and mixed turbidites. In slope and base of slope
settings, the basinal hemipelagites also intercalate numerous
shallow-derived carbonate breccias and bioclastic turbidites
(Baceta, 1996). These resedimented deposits usually occur
in the form of gully systems and apron-like accumulations
(Baceta, 1996; Baceta et al., 2004; Pujalte et al., 1993;
Robador, 2005). Landwards, the carbonate platform
succession interfingers with alluvial-lacustrine coastal
plain sediments. These non-marine deposits, extensively
developed in the south-central Pyrenees, are classically
known in the literature as the “Garumnian” facies, Tremp
Formation, or Tremp Group (Cuevas, 1992; Eichenseer,
1988; López-Martínez and Peláez-Campomanes, 1999;
Rosell et al., 2001; Schmitz and Pujalte, 2003).
The long-term transgressive trend of the Paleocene
carbonate platforms was punctuated by third-order
oscillations in relative sea level. The stratigraphic signature
of these sea-level changes is a suite of regionally extensive
Depositional Sequences (DS), which form the building
blocks of the platform-to-basin Paleocene succession (Fig.
2). The five DS recognized have been coded according to
their estimated chronostratigraphic range. Their features fit
reasonably well with current models established for landattached carbonate platforms sensu Wright and Burchette
(1996), although their character and stacking pattern vary
significantly depending on the carbonate sediment types
involved, the presence of siliciclastic sediments and the
platform depositional profile (ramp or rimmed shelf) in each
time slice (Baceta et al., 2004, 2011). In general, Paleocene
DS consist mostly of transgressive and highstand system
tracts and contain poorly developed lowstand system tracts.
Their boundaries correspond to discontinuity surfaces with
numerous features of subaerial exposure, clear evidence
of drops in sea level. The most prominent and longest
lasting of these lowstands took place during the DanianSelandian boundary and caused the complete exposure
of the Danian platform succession and the formation of a
complex paleokarst system for a period of about 2.5m.y.
(Baceta et al., 2001, 2007). The establishment and
platform-to-basin correlation of the Paleocene DS solved
many of the incongruences and misinterpretations in the
numerous formal and informal lithostratigraphic units
established within the Pyrenean Paleocene stratigraphic
record (Garrido-Megías and Rios, 1972).

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J. Serra-Kiel et al.

Biostratigraphic studies contributed to basin-wide
correlations and the detailed chronostratigraphic calibration
of the Paleocene DS. These were accurately dated with
planktic foraminifers and calcareous nannofossils, which
abound in slope hemipelagites and some outer platform
deposits. In addition, most platform deposits contain
shallow benthic foraminifers, which occur in great quantity
and can be easily recognized in the field. They are,

Paleocene LBF from the Pyrenean Basin

therefore, the best option for dating Paleogene shallowwater platform deposits, particularly in isolated outcrops
and sections. The correlation chart in Figure 2 summarizes
the relationship between planktonic foraminifera and
calcareous nannofossil zones, the Geomagnetic Polarity
Time Scale (GPTS) and third-order DS of the Paleoceneearly Eocene of the Pyrenean basin. This correlation chart
is primarily based on reference sections from the most

FIGURE 2. Bio-chronostratigraphic framework of the Pyrenean Paleocene-lower Ilerdian platform-to-basin succession, showing depositional
sequences and main facies (modified from Baceta et al., 2004, 2011). The Geologic Time Scale used is in accordance with the 2017 International
Chronostratigraphic Chart (Cohen et al., 2013) and Vandenberghe et al. (2012) and the Paleocene stage subdivision in accordance with Global
Stratotype Section and Points (GSSPs) established in the Zumaia reference section (Schmitz et al., 2011).

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representative inner platform-to-coastal plain settings
(Tremp, Campo), the platform margin (Lizarraga pass)
and base of slope (Aixola-Ermua, Monte Urko), all
correlated with the reference basinal section of Zumaia.
This chronostratigraphic framework incorporates the
recently adopted formal position of the Danian-Selandian,
Selandian-Thanetian and Paleocene-Eocene boundaries
(Schmitz et al., 2011).

correlation of the larger foraminiferal zones (SBZ) with
those of the calcareous nannoplankton (NP CNP) and
,
planktonic foraminifera (P).

Description of the stratigraphic sections

Serraduy section

For this work, we have selected a series of stratigraphic
sections that contain larger foraminifera ranging from
transitional continental-marine realms, to inner platform,
middle platform, the platform margin, the base of slope
and hemipelagic environments. For the correlation
among the different taxa of biostratigraphic use, special
attention has been paid to stratigraphic sections with
intercalations of deposits with calcareous plankton of
biostratigraphic significance and to sections of platform
slope and hemipelagic environments with calciturbidites
that contain larger foraminifera that can be considered
penecontemporaneous to planktonic foraminifera and
calcareous nannoplankton.

Serraduy section (Fig. 4) was measured on the west and
east banks of the Isabena River and adjacent hillslopes,
all located some 500-800m to the north of the village of
Serraduy.

The profiles selected run east-northwest across
the Paleocene basin axis (Fig. 1C) and permit precise

The description of the studied sections presented below
takes as reference the Paleocene stratigraphic units defined
in Figure 2. In all stratigraphic sections, the lithology is
represented according to the legend set out in Figure 3.

UTM coordinates (ETRS89, H31) are X: 299.759, Y
:
4.688.981 (bottom) and X: 300.261, Y 4.688.350 (top).
:
The base of the section is marine sandy limestone of the
Arén Formation (Fm.) (Mey et al., 1968), Maastrichtian
in age. The non-marine deposits encompassing the first
interval described (Ma-Da to IL-1 depositional sequences)
were referred as Garumnian facies by Leymerie (1868).
Later Garrido-Megías and Ríos (1972) detailed the
stratigraphy of this part of the section separating the
continental deposits (their Laspún Fm.) from the marine

FIGURE 3. Lithological legend for Figures 5-13.

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intercalations found in the following depositional sequences
(Se/Th-1 and Th-2 DS), which build up the Navarri Fm.
Finally, the summit deposits (IL-1 depositional sequence),
are included in the Serraduy Fm. (Cuevas-Gozalo et al.,
1985). The stratigraphic succession can be divided into the
following DS:
Ma-Da and Da-1 DS. The boundary between these
two DS is difficult to determine. They constitute the lower
half of the section, represented by 100m of non-marine
red clays with sandstone and silt intercalations including
pedogenetic horizons.
Da-2 DS. It is represented by lacustrine limestone
capped by two well-developed pedogenetic horizons
corresponding to the Colmenar-Tremp horizon (Eichenseer,
1988).
Se/Th-1 DS. The basal limit of this sequence is the
top of the pedogenetic horizon of the previous DS. This
unit is made up of continental red and variegated clays and
includes a marine intercalation in its upper part. This level
is composed of a conglomeratic layer sandwiched between
two shallow marine limestone beds with foraminifera. The
uppermost part of the unit is again composed of red clays
with nodular calcareous concretions. The occurrence of I.
sinjarica, G. primaeva, K. aquitanica and C. carniolica
in this stratigraphic interval characterizes SBZ 3.
Th-2 DS. This depositional sequence starts with a
1m-thick sandy conglomeratic bed, followed by 10m of
sandy limestone with abundant marine fossils, including
oysters, corals and abundant larger foraminifera, followed
by sandstone. The upper part of this DS consists of
greenish-red bioturbated continental clays. The occurrence
of D. garumnensis and G. levis in this DS is characteristic
of SBZ 4.
The lower undifferentiated red clay unit (DS Ma-Da and
Da-1) and the depositional sequences Da-2, Se/Th-1 and
Th-2 have been referred to as the Laspún Fm. by GarridoMegías and Ríos (1972).
IL-1 DS. The bottom of this unit is formed by an
incised channel with a conglomeratic infill overlain by a
6m-thick layer of calcareous sandstone and red clays. The
remaining of the sequence is composed of two limestone
intervals separated by a unit built up of unconsolidated rock,
generally covered, consisting of marlstone and calcareous
sandstone with plant remains, which indicate regressive
conditions. The first of these two calcareous beds is a
bioclastic, shallow marine limestone with very abundant
Alveolina, that has been interpreted as as a giant sandwave.
The occurrence in this DS of A. vredenburgi vredenburgi
and A. globula indicates SBZ 5 in the lower part of IL-1,

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whereas in its upper part A. vredenburgi tumida and A.
ellipsoidalis are indicative of SBZ 6. These deposits are
included in the Serraduy Fm. (Cuevas-Gozalo et al., 1985).
Campo section
Campo section (Fig. 5) is located in the Ésera river
gorge, south of the town of Campo. The section begins
at km 4 along the N-260 road, following the path of the
ancient, abandoned road, and crossing the Ésera Bridge
towards the town of Ainsa.
UTM coordinates (ETRS89, H31) are X: 285.738, Y
:
4.696.911 (bottom) and X: 285.660, Y 4.696.180 (top).
:
This section exposes a relatively thick succession of
Paleocene to lower Ilerdian platform carbonates, arranged
in up to six depositional sequences (Fig. 5). The underlying
Maastrichtian deposits are sandy limestones pertaining to
the Arén Fm. The depositional sequences Ma-Da, Da-1
and Da-2 belong to the Laspún Fm. (Garrido-Megías and
Ríos, 1972), whereas Se/Th-1 and Th-2 deposits build
up the Navarri Fm. (Garrido-Megías and Ríos, 1972).
The continental deposits from the top of the Arén Fm. to
the base of the Se/Th-1 sequence were characterized as
Garumnian facies by Leymerie (1868). Ma-Da DS. This is
a heterolithic unit composed of sandstones and marls in
its lower part, a sandstone unit with rudist debris in the
middle part, and an upper part build up of dolomicrite and
red clays.
Da-1 DS. It is made up of secondary dolomites with
calcareous algae and microfossil ghosts.
Da-2 DS. It is formed by white dolomicrites in its
lower part, capped by some beds of oolitic limestones of
lacustrine facies. The upper part begins with a collapse
breccia composed of honeycomb-like dolomite followed by
red and variegated clays and lacustrine limestone. Its upper
limit is marked by a widespread paleosol catena, called
the Colmenar-Tremp Horizon (Eichenseer, 1988). Se/Th-1
DS. The basal levels are composed of red-grey claystones
and dolomites. The main body of this unit is composed
of bioclastic, very fossiliferous limestones, deposited in
lagoon/inner shelf environments. The regressive upper
part of the section includes dolomites accumulated in
supratidal marsh environments including an intercalation
of continental clays with vertebrate remains (Tambareau
et al., 1992a). The larger foraminifera G. primaeva,
V. anatolica, C. rajkae, C. carniolica, K. aquitanica,
D. praegarumnensis, M. yvettae, M. primitivus and C.
ovoidea characterize SBZ 3.
Th-2 Ds. This is a transgressive unit composed mainly
of very fossiliferous bioclastic limestones. Its central part
has been interpreted as the deepest facies association of

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J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

FIGURE 4. Stratigraphic succession and distribution of larger foraminifera in the Serraduy section.

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the entire Paleocene interval, and is represented by marls
and marly limestones with autochthonous assemblages of
calcareous nannofossils (Orue-Etxebarria et al. 2001). The
middle and upper parts of this sequence is characterized
by sandy, bioclastic and cross-bedded bioclastic limestones.
In the uppermost part of the sequence, there are reefal

boundstones. The top of this DS is a 5m-thick layer
of mudstones with exhibits pedogenetic alteration and
abundant Microcodium, indicating subaerial exposition.
The presence in this unit of G. levis, D. garumnensis,
A. yvettae and A. azilensis characterizes SBZ 4. The
calcareous nannofossil assemblages recorded in the nine

FIGURE 5. Stratigraphic succession and distribution of larger foraminifera in the Campo section.

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samples collected in the upper part of Th-2 are indicative
of Zone NP9 of Martini (1971) or Zone CNP11 of Agnini
et al. (2014).

by bioclastic limestone rich in alveolinids. The occurrence
N. aff. minervensis and A.v. vredenburgi characterizes
SBZ 5.

IL-1 DS. Thi unit begins with continental clays and
sandstones, where the geochemical signal of the CIE of
the Paleocene-Eocene thermal maximum has been located
(Pujalte et al., 2009b; Schmitz and Pujalte, 2003). The
upper part of the unit is made up of bioclastic limestone
with Alveolina including A.v. vredenburgi, which
characterizes SBZ 5.

Garralda section
Garralda section (Fig. 7) was measured along a path that
leaves the village of Garralda towards the north. It begins at
the base of an abandoned limestone quarry and continues
northward after a small water reservoir.

Tena section

UTM coordinates (ETRS89, H30) are X: 639.894, Y
:
4. 756.777 (bottom) and X: 639.879, Y 4. 756.928 (top).
:

Tena section (Fig. 6) is located in the Gállego river
gorge at the entrance of the Tena valley, north of the town
of Biescas along the A-136 road in the vicinity of the
kilometer point 4 and along the access path to the fort and
hermitage of Santa Elena.

The rocks underlying the Paleocene succession are marly
limestones belonging to the “Calcischistes à Navarelles”
(Souquet, 1967) of Maastrichtian age. The DS Da-2
constitutes the Abaurrea Fm., while the DS Se/Th-1 and
Th-2 and IL-1 belong to the Berrendi Fm. (Robador, 2008).

UTM coordinates (ETRS89, H30) are X: 719.322, Y
:
4.726.687 (bottom) and X: 719.439, Y 4.726.505 (top).
:

Da-2 DS. This unit is represented at its lowest part by
a 1.5m-thick breccia composed of clasts made of limestone
and marly clay chips. Above, the succession is composed
of a 40m-thick, monotonous succession up of coarsening
upwards bioclastic limestones with calcareous algae
(rhodophyte) bioclasts. The occurrence in this unit of M.
globularis, O. moorkensii and E. dienii is characteristic
of SBZ 2.

The rocks underlying the Paleocene succession belong
to the Marboré Fm. (Souquet, 1967) ands are mostly
composed of sandstone and marlstone. Both Da-1 and Da-2
depositional sequences are referred to in the literature as
the Salarons Fm. (Van de Velde, 1967). The depositional
sequences Se/Th-1, Th-2 and IL-1 belong to the Gallinera
Fm. (Van de Velde, 1967).
Da-1 DS. This unit is composed of dolomites and
shallow marine limestones capped with intraformational
breccia at top. The occurrence of B. hanseni at top is
characteristic of SBZ 1.
Da-2 DS. This unit is characterized by shallow marine
limestone with algal laminations, dolomites and shallow
marine limestones at the top. Unfortunately, the absence of
the larger foraminifera does not alow for a determination of
the biostratigraphic range of this DS.
Se/Th-1 DS. The basal levels of this depositional
sequence are characterized by sandstone and shallow
sandy limestone, followed by algal boundstones and topped
by shallow marine limestones. The occurrence of G.
primaeva, V. anatolica, F. alavensis, C. carniolica, “O”.
heberti and D. seunesi characterizes SBZ 3.

Se/Th-1 DS. This unit is composed in its lower part
by sandy, fine grained limestones, which overlies Da-2
deposits with a sharp, erosive contact. The upper half of
the sequence is made of bioclastic limestones, where V.
anatolica, K. aquitanica and M. yvettae where found and
indicate SBZ 3.
Th-2 DS. This unit is composed in its basal levels
by sandy limestones which abruptly overlie the bioclastic
limestones of the previous sequence followed by bioclastic
limestones with calcareous algae and corals.
IL-1 DS. This DS is made of alternating marls and
marly limestones with abundant planktonic foraminifera.
Due to their incompetent nature, their outcrops are very
scarce, with its lower boundary being usually covered.
Urrobi section

Th-2 S. This unit is composed of very fossiliferous
shallow marine sandy limestones. The occurrence of G.
levis, A. yvettae and A. azilensis characterizes SBZ 4.

Urrobi section (Fig. 8) was measured in year 1994 on
the left bank of the Urrobi river south of the village of
Nagore; however, it is no longer accessible as the area has
been flooded by the Itoiz reservoir.

IL-1 DS. The bottom of this unit is represented by
bioclastic sandy silt with abundant nummulitids followed

UTM coordinates (ETRS89, H30) are X: 633.448; Y
:
4.743.460 (bottom) and X: 633.487; Y 4. 743.057 (top).
:

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Da-2 DS. This unit is represented by primary dolomites
in its lower part and shallow marine limestones including
bioclastic beds in the upper half. Unfortunately, the absence
of larger foraminifera in this stratigraphic interval does not
allow determining its biostratigraphic range.

Da-1 DS. This unit is composed of shallow-marine
dolomites with occasional interbedding of limestones. The
presence of B. hanseni indicates SBZ 1 in the upper part
of this unit.

Se/Th-1 DS. The basal levels are characterized by
shallow marine sandy limestones, followed by alternating
marlstones and sandy limestones. The top of the sequence
is formed by boundstone and bioclastic marine limestone.

“O.”

The rocks underlying the Paleocene succession
belong to the Marboré Fm. represented by sandstones and
marlstones. Both Da-1 and Da-2 depositional sequences
belong to the Salarons Fm., while the DS Se/Th-1, Th-2
and IL-1 DS belong to the Gallinera Fm.

FIGURE 6. Stratigraphic succession and distribution of larger foraminifera in the Tena section.

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Paleocene LBF from the Pyrenean Basin

FIGURE 7. Stratigraphic succession and distribution of larger foraminifera in the Garralda section.

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The occurrence of “O”. heberti and D. seunesi indicates
SBZ 3.
Th-2 DS. This unit is composed of marlstone and very
fossiliferous sandy limestones. The presence of A. yvettae
and A. azilensis indicates SBZ 4.
IL-1 DS. The bottom of this sequence is formed by
bioclastic sandy silt with abundant nummulitids where
the geochemical signal of the CIE is located according

to Pujalte et al. (2003). These rocks are followed by a
bioclastic limestone rich in nummulitids, with N. aff.
minervensis indicating SBZ 5.
Andia section
Andia section (Fig. 9) was logged on the edge of the
plateau located approximately 1.5km to the east of the
Lizarraga mountain pass. It is provided here as subsidiary
section of the Lizarraga section, described below. This

FIGURE 8. Stratigraphic succession and distribution of larger foraminifera in the Urrobi section.

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section is relevant in the Western Pyrenees as it is the only
including Selandian deposits in shallow water facies, most
of them relatively rich in larger foraminifera.
UTM coordinates (ETRS89, H30) are X: 582.279, Y
:
4.746.093 (bottom) and X: 581.933, Y 4.745.805 (top).
:
Se/Th-1 DS
This sequence lies unconformably on upper
Danian “Lizarraga reef complex” and is overlain by
unnamed Ilerdian deposits. The succession is made up
of marlstones and marly limestones interbedded with
several m-thick intervals of bioclastic limestones and
very locally coralgal boundstones (Baceta et al., 2005).
The stratigraphic interval logged is laterally equivalent
to the informal unit known as Usabide reef complex
(Baceta et al., 2005). The bioclastic limestones comprise
a varied association of foraminifera indicative of the
SBZ 3 biozone, namely, G. primaeva, V. anatolica, C.
rajkae, M. primitivus, “O”. heberti and D. seunesi.
According to G. Bernaola in Baceta et al. (2005), the
logged section encompasses the NP5, NP6, NP7 and
NP8 calcareous nannofossil zones of Martini (1971).
The revision of these data allowed us to recognize in
the same interval the CNP7, CNP8, CNP9 and CNP10
calcareous nannofossil zones of Agnini et al. (2014).

Lizarraga section
Lizarraga section (Fig. 10) is located along the NA120 road from Etxarri-Aranatz to Estella/Lizarra across
Lizarraga mountain pass, on the northern edge of the
Urbasa-Andia synclinorium.
UTM coordinates (ETRS89, H30) are X: 581.722, Y
:
4.746.147 (bottom) and X: 581.029, Y 4.745.731 (top).
:
The section consists of an up to 360m-thick Paleocene
to Early Eocene platform margin succession including
different intervals with calcareous nannofossils allowing
for an accurate age determination. The Paleocene overlies
conformably the Maastrichtian deposits (Puerto de
Olazagutia Fm., Amiot, 1982). The Da-2 DS is informally
known as the Lizarraga reef complex, while the Se-Th-1
DS is laterally equivalent to the so-called Legumbe reef
complex and DS Th-2 to the so-called “Assilina beds”
.
(Baceta et al., 2005).
Ma-Da DS. This unit is composed of marly limestones
and fine-grained limestones lacking shallow benthic
foraminifera. The top is characterized by an irregular bed
of breccia interpreted as a slump. The NP1 or CNP1 (basal
Danian) age of this unit was established in by Baceta et al.
(2005) based on calcareous nannofossils.

FIGURE 9. Stratigraphic succession and distribution of larger foraminifera in the Andia section.

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zone. This indicates a basal hiatus between this and the
former unit, which encompasses the biozone CNP2 and at
least the lower half of CNP3.
Da-2 DS. This unit is mostly made up of near-massive
limestones (reefal boundstones) with abundant colonial
corals and calcareous algae (dasyclads, solenoporaceans and
coralline red algae). These deposits contain abundant larger
foraminifera, such as K. decastroi, H. paleocenica, H.
elongata, E. dienii and O. moorkensii, characteristic of

“O.”

Calc. nannofossils
Zones

Calc. nannofos. samples

Da-1 DS. This unit is largely composed of marlstone
and marly limestone. In the upper part, larger foraminifera
such as S.? binkhorsti and P.? cretae were found, both
without biostratigraphic interest. According to Baceta et
al. (2005), the unit belongs to the lower part of NP3 zone
with a basal hiatus spanning NP2 and probably the base of
NP3. This is confirmed with data from laterally-equivalent
deposits in the Unanua III and Baiza peak sections (see
Baceta et al. 2005). Taking into account the zonation of
Agnini et al. (2014), this interval belongs to the CNP3

FIGURE 10. Stratigraphic succession and distribution of larger foraminifera in the Andia section.

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SBZ 2. According to Baceta et al. (2005), the unit belongs
to the upper part of biozone NP3 as evidenced in laterallyequivalent deposits in the Unanua III section. The top of the
unit is marked by a prominent discontinuity showing features
of subaerial exposure, which is regionally known as the midPaleocene unconformity (Baceta et al., 2004, 2007).
Se/Th-1 DS. This is an up to 20-25m-thick and
unit made up of marly limestones evolving vertically to
bioclastic limestones. The latter are rich in larger
foraminifera such as “O.” heberti, D. seunesi and lessabundant G. primaeva, C. rajkae and M. primitivus, all
representative of SBZ 3. This DS is laterally equivalent to
the so-called Legumbe reef complex (Baceta et al., 2005).
In the lower marly interval of the Se/Th-1 DS, Baceta et
al. (2005) documented calcareous nannofossil assemblages
representative of NP8 or CNP10 zones, and thus indicative
of a Thanetian age.
Th-2 DS. This is a comparatively thin unit, mostly
composed of marlstone and marly limestones of open
platform deposits with abundant planktonic fossils,
laterally-equivalent to the upper Thanetian “Assilina beds”
(Baceta et al., 2005). Calcareous nannofossils are of late
Thanetian age, specifically belonging to Zones NP9 or
CNP11 (Baceta et al., 2005).
IL-1 DS. This unit is composed of marly limestone,
marls and limestone. The NP10 or CNE2 (Eocene) age of
this unit was established by Baceta et al. (2005).
Urbasa Pass section
Urbasa Pas section (Fig. 11) is located on the northern
flank of the Urbasa-Andia synclinorium, along the NA-718
road from the Urbasa (Olazagutia) road pass to the Bidoiza
campsite. It represents an up to 300m-thick Paleocene to
lower Eocene succession belonging to marine platform
facies lacking Selandian deposits.
UTM coordinates (ETRS89, H30) are: X: 546.182, Y
:
4.727.412 (bottom) and X: 567.787, Y 4.744.504 (top).
:
The Paleocene rocks lie on erosional contact over
the Maastrichtian Puerto de Olazagutia Fm. (Amiot,
1982). The Ma-Da DS has been informally called the
“Solenomeris limestone” while the Th-2 was called the
,
“Assilina beds” by Baceta et al. (2005). The same authors
called the deposits of IL-1 DS the “Nummulites-Alveolina
limestone”
.
Ma-Da DS. This unit is made of bioclastic limestones
rich in the encrusting foraminifer Solenomeris sp., red
algae (rhodophytes), bryozoans and B. hanseni indicating
the SBZ 1 biozone.

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Da-1 DS. This unit is composed of stratified to massive
bioclastic limestone and reefal boundstones culminating
with cross-bedded deposits. Most of these facies are
affected by intense but patchy secondary dolomitization.
This stratigraphic interval was informally defined as the
Santa Marina reef complex by Baceta et al. (2005). The
lesser-dolomitized beds contain M. globularis and O.
moorkensii indicating SBZ 2 (lower part).
Da-2 DS. It is made in the lower half of cyclicallyordered bioclastic lagoonal limestones and marlstones,
with patchy secondary dolomitization. These beds,
interpreted as belonging to lagoonal facies, are laterally
equivalent to the Lizarraga reef complex (Baceta et al.,
2005). The bioclastic limestone beds contain K. decastroi,
H. paleocenica, H. elongata and M. globularis, indicating
SBZ 2 (upper part).
Se/Th-1 DS. The unit begins with a marlstonedominated interval that vertically evolves into bioclastic
and massive red algae-coral boundstones overlain by
bioclastic limestones, eventually with cross bedding and
sandy intercalations. These beds are laterally equivalent to
the Legumbe reef complex (Baceta et al., 2005). The DS
contains a rich association of larger foraminifera, including
G. primaeva, M. juliettae, M. primitivus, “O.” heberti, R.
soldadensis and D. seunesi, which belong to SBZ 3.
Th-2 DS. It is composed of marlstone and thin beds of
bioclastic limestones. These contain numerous A. azilensis
indicating SBZ 4.
IL-1 DS. It is characterized by bioclastic limestones
with A. v. vredenburgi and N. gamardensis, indicating
SBZ 5.
Leortza section
Leortza section (Fig. 12) is located along the A3114
road, in the river gorge located between the localities of
Leortza/Elortza and Cicujano/Zekuiano, on the northern
flank of the Maestu diapir. It encompasses a complete
succession of shallow marine Maastrichtian to lowermost
Eocene carbonates and siliciclastic rocks. However, the
present study focuses on the upper Paleocene interval (Se/
Th-1 and lower part of the Th-2 sequences).
UTM coordinates (ETRS89, H30) are X: 545.651, Y
:
4.733.233 (bottom) and X: 545.722, Y 4.733.446 (top).
:
The stratigraphic interval represented by DS Se/Th-1
was informally called the “Legumbe reef complex” while
,
the deposits of Th-2 DS were called the “Assilina beds”
(Baceta et al., 2005).

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V. anatolica, C. rajkae, F. alavensis, M. yvettae,
M. juliettae, “O.” heberti and D. seunesi among
others indicates SBZ 3 for this stratigraphic interval.
Additionally, it includes the type-locality beds of
Fallotella alavensis.

“O.”

v. vredenburgi

Th-2 DS. This unit consists of bioclastic sandy silts
with intercalations of bioclastic limestones rich in A.
azilensis indicating SBZ 4.

patalaensis

Se/Th-1 DS. This unit lies unconformably on
dolomitized and karstified Danian deposits of the San
Justi Fm. (Mangin, 1960) and begins with a thin and
discontinuous bed of Qz-rich microconglomerates
and sandy limestones. The bulk of the sequence
consists of bioclastic limestones comprising a
prominent interval, rich in red algae, bryozoans and
corals, while stratified marlstones and dolomites
occur atop. The presence of the larger foraminifera

FIGURE 11. Stratigraphic succession and distribution of larger foraminifera in the Urbasa Pass section.

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Korres section
The section (Fig. 13) is located in some cliff outcrops 300m
west of the town of Korres, close to the A-3138 road to Durruma
Kampezu. The section comprises a complete succession of
upper Paleocene to lower Eocene deposits (Th-2 and Il-1).

Th-2 DS. The lower part of this unit is formed by
sandy limestones with cross stratification, grey silt and sandy
marlstone that vertically evolve into bioclastic limestones. This
stratigraphic interval has been informally named the “Assilina

“O.”

UTM coordinates (ETRS89, H30) are X: 546.182; Y
:
4.727.412 (bottom) and X: 546.131; Y 4.727.608 (top).
:

The studied units overly a lower-middle Thanetian
succession equivalent to the Se/TH-1 sequence at Leortza.
The interval corresponding to the DS IL-1 was informally
called the Ilerdian Alveolina limestone (Baceta, 1996).

FIGURE 12. Stratigraphic succession and distribution of larger foraminifera in the Leortza section.

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beds” (Baceta et al., 2005). The top is a prominent paleokarst
surface that defines the Paleocene-Eocene boundary (Baceta,
1996). The presence in this stratigraphic interval of G. levis,
H. lukasi, D. garumnensis and A. yvettae indicates the SBZ
4. The sample K 2 located in this DS is the type-locality of the
new species: A. korresensis and V. bacetai.

(Aixola-Egoarbitza
and
Er mua)
located
respectively on the souther n and nor ther n f lanks
of the Egoarbitza syncline (SE tip of the Biscay
sinclinorium). Both are accessible from the town
of Er mua, along the local GI-121 road to the
Aixola dam and reservoir.

IL-1 DS. It consists of a basal 6-7m-thick interval of
unfossiliferous marlstone and sandy limestone with cross
stratification overlain by bioclastic limestones including A.
aramea aramea and A. v. vredenburgi which indicates SBZ 5.

The Aixola-Egoarbitza section is located just north
of the dam, with UTM coordinates (ETRS89, H30)
X: 539.896, Y 4.779.091 (bottom) and X: 539.924, Y
:
:
4.779.244 (top).

Aixola-Ermua section

The Ermua section is located closer to the Ermua town,
on the banks of the local road, with UTM coordinates
(ETRS89, H30) X: 540.794, Y 4.780.716 (bottom) and X:
:
540.728, Y 4.780.681 (top).
:

Aixola-Er mua section (Fig. 14) is a composite
section from two laterally-equivalent outcrops

FIGURE 13. Stratigraphic succession and distribution of larger foraminifera in the Korres section.

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The underlying Maastrichtian deposits are marlstones
with turbidite intercalations of the Urko Fm. (Mathey,
1982). In both outcrops, the Paleocene succession is a
complex series of base-of-slope and basinal deposits,
approaching 122m in composite thickness, arranged in five
DS. The deposits of the Da-1 and Da-2 DS belong to the
Aitzgorri Fm. (Bernaola et al., 2009), while those of the
Se/Th-1 and Th-2 DS belong to the Itzurun Fm. (Baceta et
al., 2005).
Ma-Da DS. This unit is only well-exposed in some
accessory outcrops to the east of the main Aixola outcrop,
where it comprises 14m of the upper Maastrichtian deposits,
the K/Pg boundary layer and 6m of hemipelagic limestone
with thin calcareous turbidites of earliest Danian age.
Da-1, Da-2, Se/Th-1 and Th-2 DS. All these units
are bounded by discontinuity surfaces and internally consist
of calcareous breccias deposits overlain by hemipelagic
limestones and marlstones with interbedded platform and
slope-derived calciturbidites and/or siliciclastic turbidites.
The occurrence of E. dienii and M. globularis in the
Da-2 DS indicates SBZ 2, while the Se/Th-1 DS contains G.
primaeva, F. alavensis, R. soldadensis and “O.” heberti,
indicating SBZ 3. The SBZ 4 biozone is characterized by
D. garumnensis, E. subsphaerica, A. yvettae and A.
azilensis observed in the Th-2 DS.
Calcareous nannofossil data from this study, allow the
identification of the Selandian-Thanetian boundary in an
interval a few meters above the NP5-NP6 zone boundary.
This interval also records the first occurrence of “O.”
heberti (SBZ 3). The calcareous nannofossil assemblages
studied here from the Da-2 and Se/Th-1 DS correspond
to NP4, NP5, NP6, NP7 and NP8 of Martini (1971) and
CNP6, CNP7, CNP8, CNP9 and CNP10 zones of Agnini et
al. (2014) (Figure 14).
The upper part of the section (Th-2 DS and IL-1 DS)
was previously studied by Monechi in Orue-Etxebarria
et al. (1996), who attributed them to the calcareous
nannoplancton Zones NP9 and NP10 respectively,
which correspond to CNP11 and CNE1 of Agnini et al.
(2014). The zonation of this section based on planktonic
foraminifera was established by X. Orue-Etxebarria in
Pujalte et al. (1993).
IL-1 DS. The base of this sequence is an erosional
discontinuity marking an abrupt change to clay-rich
facies with turbidites of mixed composition containing
larger foraminifera indicative of SBZ 5, such as N. cf.
gamardensis. The geochemical signal of the CIE marking
the Paleocene-Eocene boundary is located in the base of
this DS according to Pujalte et al. (2009a, b).

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Urko section
Urko section (Fig. 15) is located on the northern flank
of the Biscay sinclinorium, on the eastern hillside of Urko
peak, some 3km north of the towns of Eibar and Ermua
(Gipuzkoa province). The section is accessible along a path
from the Ixua road pass, onthe GI-3950 road.
UTM coordinates (ETRS89, H30) are X: 541.754, Y
:
4.783.428 (bottom) and X: 541.722, Y 4.783.390 (top).
:
Overlying the Monte Urko Fm. composed of typically
reddish-purple hemipelagic rocks and above the K/Pg
boundary layer, there are grey-pinkish limestones and
marlstones with numerous intercalations of platformderived calciturbidites belonging to the Danian stage
(Baceta, 1996). This Danian succession is equivalent to
the Aitzgorri limestone Fm. exposed in the type locality
of Zumaia (Bernaola et al., 2009), and as there it can be
subdivided into three intervals corresponding respectively
to the Ma-Da, Da-1 and Da-2 DS. Their boundaries are
conformity surfaces without recognizable stratigraphic
hiatus.
The absence of larger foraminifera in most platformderived turbidites of the Ma-Da DS and the lower part
of the Da-1 DS prevent a precise characterization of the
SBZ1-SBZ2 boundary. SBZ 2 is indicated by the larger
foraminifera K. decastroi, E. dienii, O. moorkensii and P.
eos from the middle part of the Da-1 to the Da-2 DS.
Calcareous nannofossils of the 29 samples collected
in the Danian of Urko section allowed the identification
of four calcareous nannofossil zones defined by Martini
(1971) for the Danian and the seven zones defined by Agnini
et al. (2014) for the same interval (Fig. 5). Following the
amendments introduced by Perch-Nielsen (1981, 1985), we
have used the first occurrences of Cruciplacolithus primus
large and Cruciplacolithus edwarsii to define the base of
zones NP2 and NP3. Ellipsolithus macellus, whose first
occurrence marks the base of NP4, is very scarce and it
has a discontinuous distribution in the upper part of the
section.
Systematic Palaeontology (J. Serra-Kiel and V.
Vicedo)
Phylum: FORAMINIFERA d’orbigny
Class Tubothalamea pawlowski et al.,
Order Miliolida (delage and hérouard)
Superfamily Milioloidea ehrenberg
Family Haurenidae schwager
Subfamily Miliolinellinae vella
Genus Idalina Schlumberger and munier chalmas
Type species: Idalina antiqua schlumberger and
munier-chalmas, 1884

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“O.”

NP4

CNP6

FIGURE 14. Stratigraphic succession and distribution of larger foraminifera in the Aixola-Ermua section. Red points indicate reworked specimens.

P1a+b

P1c

P3

CNP
10

11

CNE

NP 2
10 CNE
1

SBZ
1

J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

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J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

FIGURE 15. Stratigraphic succession and distribution of larger foraminifera in the Monte Urko section.

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J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

Idalina sinjarica grimsdale, 1952
Figs. 16A-16D

Kayseriella decastroi sirel, 1998
Fig. 16E-16H

1952 Idalina sinjarica. Grimsdale, p. 230, pl. 1, fig.
1-13; pl. 12, fig. 1,2; pl. 13, fig. 2; pl. 14, fig. 1
1974 Idalina sinjarica grimsdale. Drobne, p. 166-167;
pl. 1, fig. 1-2, 4-13
1998 Idalina sinjarica grimsdale. Sirel, p. 55-56; pl.
17, fig. 1-23
2001 Idalina sinjarica grimsdale. Özgen and Akyazi,
pl. 1, fig. 12-13; pl. 2, fig. 1
2008 Idalina sinjarica grimsdale. Pignatti et al., pl. 5,
fig. 4; pl. 7, fig. 11
2010 Idalina sinjarica grimsdale. Di Carlo et al., p.
54-55; pl. 4, figs. 1-19
2015 Idalina sinjarica grimsdale. Sirel, pl. 19; figs.
1-23

1988 “Scandonea” Drobne et al., pl. 25, fig. 7
.
1998 Kayseriella decastroi Sirel, p. 57-58; pl. 16, fig.
19-23; pl. 20, fig. 1-11; pl. 21, fig. 1-12; pl. 22, fig. 1-15
1999 Kayseriella decastroi Sirel, p. 128; pl. 1, fig.
1-11; pl. 2 fig. 1-12; pl. 3, 1-15
2001 Kayseriella decastroi sirel, Ogorelec et al., pl.
9, fig. 4-7
2004 Kayseriella decastroi sirel, p. 60-61; pl. 55,
figs. 1-11; pl. 56, figs. 1-12; pl. 57, figs. 1-15
2009 Kayseriella decastroi sirel, pl. 2, figs. 1-7
2010 Kayseriella decastroi sirel, Di Carlo et al., p.
54; pl. 1, fig. 2
2012 Kayseriella decastroi sirel, pl. 1, figs. 19-23
2015 Kayseriella decastroi sirel, pl. 1, figs. 19-23; pl.
6; figs. 1-12
2018 Kayseriella decastroi sirel, pl. 43, figs. 1-12

Material. Sample Se 1 from Serraduy section (Fig.
4); samples Cm 5 and Cm 8 from Campo section (Fig. 5);
samples Te 3 and Te 4 from Tena section (Fig. 6); sample L
27 from Lizarraga section (Fig. 10); samples Le 7 and Le
8 from Leortza section (Fig. 12) and sample Coll 3 from
Collarada section (Robador, 2005, Annex p. 22-23).
Description. Test porcelaneous with ovoid
morphology and miliolid growth. Dimorphism marked.
The megalospheric forms show an elliptical outline in
longitudinal section and subcircular outline in perpendicular
section. The major diamenter for three whorls is 0.8mm and
1.25mm for four whorls. The proloculus is spherical, 140170µm in diameter, with a flexostyle followed by chambers
in quinqueloculine arrangement, becoming bilocular in
later stages of growth. The trematophore is supported by
thin pillars (Fig. 16B-16D). The undulate basal layer does
not exceed half of the chamber height (Fig. 16B-16D).
The microspheric generation has a thick basal layer and a
subcircular outline in axial section (Fig. 16).
Distribution. This species has been identified in Se/
Th-1 DS of the following sections: Serraduy section (Fig.
4); Campo section (Fig. 5); Tena section (Fig. 6); Lizarraga
section (Fig. 10) and Leortza section (Fig. 12) with the
species assemblage indicated in the aforementioned figures.
Biozone. Drobne (1974) identified this species
in rocks from Slovenia considered as belonging to the
interval from G. primaeva Zone to A. ellipsoidalis Zone.
According to Serra-Kiel et al. (1998), its biostratigraphic
range extends from SBZ 3 to SBZ 6. In this work I.
sinjarica is associated to assemblages belonging to the
SBZ 3.
Genus Kayseriella sirel
Type species: Kayseriella decastroi sirel

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

Material. Sample L 20 from Lizarraga section (Fig.
10); samples Urb 13, Urb 14, Urb 15 and Urb 18 from
Urbasa Pass section (Fig. 11); sample Ur 9 from Urko
section (Fig. 15) and samples Ain 5 and Ain 30 from
Aintzioa section (Robador, 2005, Annex, p. 64-65).
Description. Test porcelaneous with lenticular, flat
morphology. The proloculus has a diameter of 70µm
followed by 8-9 planispiral chambers (Fig. 16F H). The
,
single aperture, located at the bottom of the apertural face,
shows a peristome (Fig. 16E, H). Later stage of growth
composed of chambers arranged in an uncoiled-uniserial
pattern with cribrate apertures. Microspheric forms not
found.
Distribution. This species is present in Da-1 DS
of the Urko section (Fig. 15), , in the Da-2 DS of the
Lizarraga section (Fig. 10), and in the Urbasa Pass section
(Fig. 11), with the species assemblage indicated in the
aforementioned figures.
Biozone. Sirel (1998) found this species in Danian
rocks from Turkey. Later, Ogorelec et al. (2001) found it in
the Danian and Selandian from Slovenia. According to the
data presented herein this species can be considered as a
marker of the SBZ 2.
Family Fabulariidae ehrenberg
Genus Periloculina munier-chalmas and
schlumberger

Type species: Periloculina zitteli munier-chalmas
and schlumerger

Periloculina slovenica drobne, 1974
Fig. 16I-16L

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Paleocene LBF from the Pyrenean Basin

FIGURE 16. Paleocene Larger Foraminifera from the Pyrenean Basin. A-D: Idalina sinjarica grimsdale, 1952 A axial section, microspheric form; B
and C centered longitudinal sections, megalospheric forms; D uncentered axial section. Specimen A from sample Cm 8; B from sample L 27 and C
and D from sample Coll 3. E-H: Kayseriella decastroi sirel, 1999. E oblique section; F and H equatorial sections; G axial section. Specimen E from
sample Urb 13; F from sample Ur 9; G from sample Ain 30 and H from sample Urb 15. I-L: Periloculina slovenica drobne, 1974 I oblique section,
microspheric form; J and L oblique sections, megalospheric forms; K longitudinal section, megalospheric form. Specimens I and J from sample An
9; K from sample Cm 5 and L from sample Le 8. M-Q: Pseudolacazina donatae (drobne, 1974) M centered axial section, young microspheric form;
N centered axial section, microspheric form; subaxial section; O subaxial section; P oblique section; Q centered axial section, megalospheric form.
Specimen M from sample Coll 3; N from sample Te 4; O and P from sample Cm 5 and Q from sample Le 8. R-U: Hottingerina lukasi drobne, 1975
R and S subaxial sections; T and U oblique sections. Specimens R-T from sample K 2 and U from sample K 1. Abbreviations: pr: proloculus, th:
trematophore, fl: flexostyle, bl: basal layer, b: beam

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J. Serra-Kiel et al.

1974 Periloculina slovenica. Drobne, p. 173-175, textfig 12c, d; pl. 11, fig. 1-10; pl. 12, fig. 1
1984 Periloculina slovenica drobne. Drobne, p. 1419; pl. 4, fig. 1-15; pl. 5, fig. 1-7; pl. 6, fig. 1-9
2001 Periloculina slovenica drobne. Özgen and
Akyazi, pl. 1, fig. 14; pl. 2, fig. 1
2010 Periloculina slovenica drobne. Di Carlo et al., p.
55-56; pl. 4, figs. 20-27, 29-33.
Material. Samples Cm 5 and Cm 8 from Campo
section (Fig. 5); samples Te 3 and Te 4 from Tena section
(Fig. 6); samples An 7, An 9 and An 10 from Andia
section (Fig. 9); sample L 26 from Lizarraga section (Fig.
10) samples Le 7 and Le 8 from Leortza section (Fig.
12) and sample Coll 3 from Collarada section (Robador,
2005, Annex, p. 22-23).
Description. The test is porcelaneous with ovoid
morphology and miliolid growth. The dimorphism is strong.
The megalospheric generation begins with a flexostylic
proloculus, of around 150µm in diameter, followed by
chambers in triloculine arrangement (Fig. 16K). The basal
layer forms longitudinal ridges, occasionally reaching the
chamber roof (Fig. 16I-J, L). The microspheric form has
a longitudinal length of 2.4mm and an axial diameter of
about 1.75mm.
Distribution. This species is found in Se/Th-1 DS of
the Campo section (Fig. 5), Tena section (Fig. 6), Andia
section (Fig. 9), Lizarraga section (Fig. 10) and Leortza
section (Fig. 12), with the species assemblage indicated in
the aforementioned figures.

Paleocene LBF from the Pyrenean Basin

Material. Samples Cm 5 and Cm 8 from Campo
section (Fig. 5); samples Te 4 and Te 5 from Tena section
(Fig. 6); sample Le 8 from Leortza section (Fig. 12) and
sample Coll 3 from Collarada section (Robador, 2005,
Annex, p. 22-23).
Description. Porcelaneous test with spherical
morphology and miliolid growth. The dimorphism is
marked. Megalospheric forms have an outline elliptical
in longitudinal section and subcircular in perpendicular
section (Fig. 16Q). The proloculus is spherical with a
diameter of 140µm followed by chambers in biloculine
arrangement. The basal layer forms numerous
longitudinal partitions subdividing chambers into
chamberlets (Fig. 16M, N, Q). Microspheric forms not
found.
Distribution. This species is found in Se/Th-1 DS
of the Campo section (Fig. 5), Tena section (Fig. 6) and
Leortza section (Fig. 12), with the species assemblage
indicated in the aforementioned figures.
Biozone. Drobne (1974) found this species in the
G. primaeva Zone and in the A. ellipsoidalis Zone in
rocks from Slovenia. Drobne in Serra-Kiel et al. (1998)
extended the biostratigraphic range from SBZ 4 to SBZ
6. In the material studied P. donatae is associated with
species typical of the SBZ 3. Consequently, we extend its
biostratigraphic range from SBZ 3 to SBZ 6.
Family Soritidae Ehrenberg
Subfamily Praerhapydioninae hamaoui and
fourcade

Biozone. Drobne (1974) identified this species in
Thanetian rocks (G. primaeva Zone) from Slovenia. As
pointed out by Serra-Kiel et al. (1998) its biostratigraphic
range is SBZ 3. This species can be considered as a marker
for SBZ 3.
Genus Pseudolacazina caus
Type species: Pseudolacazina hottingeri caus
Pseudolacazina donatae (drobne, 1974)
Fig. 16M-Q
1974 Fabularia donatae Drobne, p. 167-169; text fig.
9, a-b; pl. 2, fig. 1-6; pl. 3, fig. 1-3 ; pl. 4, fig. 1-9,
7-21; pl. 9, fig. 1; pl. 13, fig. 1; pl. 14, fig. 1; pl. 15,
fig. 1-2
1998 Pseudolacazina donatae (drobne). Sirel, p. 6162; pl. 27, fig. 1-15
2008 Pseudolacazina donatae (drobne). Pignatti et
al. pl. 7, fig. 12
2009 Pseudolacazina donatae (drobne). Sirel, pl. 3,
figs. 9-13

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

Genus Haymanella sirel
Type species: Haymanella paleocenica sirel
Haymanella paleocenica sirel, 1998
Fig. 17A-M
1998 Haymanella paleocenica Sirel, p. 36-37; pl. 2,
fig. 1-18; pl. 3, figs. 1-12
1999 Haymanella paleocenica sp., Sirel, p. 124, 126;
pl. 4, figs. 1-18; pl. 5, figs. 1-13, 2001 Haymanella
paleocenica Sirel. Ogorelec et al. pl. 9, figs. 1-3
2004 Haymanella paleocenica. Sirel, p. 66; pl. 61,
figs. 1, 3, 5-9, 11, 13, 15-18; pl. 67, figs. 1, 5-9, 11, 12
2010 Haymanella paleocenica Sirel, Di Carlo et al. p.
62-63; pl. 1, figs. 5, 6; pl. 5, fig. 17
2015 Haymanella paleocenica Sirel pl. 4, figs. 7-18
2018 Haymanella paleocenica Sirel, pl. 48, figs. 7-18
Material. Samples L 5, L 10, L 14, L 15, L 16, L 17 and
L 18 from Lizarraga section (Fig. 10) and samples Urb
13, Urb 18, Urb 19 and Urb 21 from Urbasa Pass section
(Fig. 11).

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J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

Description. The test is porcelaneous with elongate,
cilindrical to high conical morphology in the adult forms.
The wall is thick and contains agglutinated grains on the
surface. The embryonic and nepionic stages have not been
adequately observed. The ephebic stage is composed of
chambers in uniserial arrangement, showing a maximum
length of 2.25mm and a diameter of 0.65mm for 8 uniserial
chambers (Fig. 17A). The sutures are slightly depressed.
The aperture is single, with a star-shaped outline (Fig.
17M) and located in interiomarginal position (Fig. 17H-J)
in planispiral chambers, while it occurs in terminal position
in the uniserial chambers (Fig. 17A-C, E-F).

and Selandian from Slovenia. According the data presented
here this species can be considered as a marker of the SBZ 2.

Distribution. This species has been identified in Da-2
DS in the Lizarraga section (Fig. 10) and in Urbasa Pass
section (Fig. 11) with the species assemblage indicated in
the aforementioned figures.

Material. Samples L 3, L 10, L 13, L 14, L 18 and L 21
from Lizarraga section (Fig. 10) and samples Urb 13, Urb
17, Urb 18, Urb 20 and Urb 21 from Urbasa Pass section
(Fig. 11).

Biozone. Sirel (1998) found this species in the Danian
rocks from Turkey Ogorelec et al. (2001) found it in the Danian
.

Description. The test is porcelaneous with elongate
morphology and thick wall. The embryonic apparatus has

Haymanella elongata sirel, 2009
Fig. 17N-17V
1999 Haymanella paleocenica. Sirel, p. 124, 126; pl.
4, figs. 4, 10, 12, 14, 18; pl. 5, figs. 1-4, 8-10
2009 Haymanella elongata. Sirel, p. 416-418; pl. 1,
figs. 11-17
2015 Haymanella elongata. Sirel, pl. 5, figs. 1-12
2018 Haymanella elongata. Sirel, pl. 49, figs. 1-12

FIGURE 17. Paleocene Larger Foraminifera from the Pyrenean Basin. A-M: Haymanella paleocenica sirel, 1999 A-G uncentered longitudinal
sections; H centered longitudinal section; I-J nepionic longitudinal sections; K-L oblique sections, note the aperture; M horizontal section, note the
star-shaped outline of the aperture. Specimens A, E, G and H from sample L 5; B and C from sample Urb 13; D from sample L 15; F from sample L
17; I and K from sample L 14; J from sample L 18; L and M from sample Urb 18. N-V: Haymanella elongata sirel, 2009 N-Q longitudinal sections;
R-T nepionic stage; U-V horizontal section, note the star-shaped outline of the aperture. Specimen N from sample Urb 13; O from sample Urb 18; P
and S from sample Urb 20; Q from sample L 18; R and T from sample L 21; U from sample L 14 and V from sample L 13. W-Z: Praerhapydionininae
indet. centered longitudinal sections. Specimen W from sample Urb 19 and X-Z from sample L 21.

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J. Serra-Kiel et al.

not been adequately observed. The nepionic stage of growth
consists of chambers planispirally arranged (Fig. 17S). The
ephebic stage shows uniserial chamber arrangement, with a
maximum longitudinal section of 1.6mm and a diameter of
0.35mm with 8 uniserial chambers (Fig. 17O). The single,
star-shaped aperture is located in interiomarginal position
in the nepionic stage (Fig. 17R) and in terminal position in
the ephebic stage (Fig. 17P Q)
,
Distribution. This species appears in Da-2 DS of the
Lizarraga section (Fig. 10) and the Urbasa Pass section
(Fig. 11) with the species assemblage indicated in the
aforementioned figures.
Biozone. sirel (2009) found H. elongata in Danian
and Selandian from Turkey. In the material studied, this
species can be considered as a marker for the SBZ 2.
Praerhapydionininae indet.
Fig. 17W-Z
Material. Samples An 3, An 4, An 8 and An 11 from
Andia section (Fig. 8); sample L 21 from Lizarraga section
(Fig. 10) and sample Urb 19 from Urbasa Pass section (Fig.
11).
Description. Porcelaneous test with a thick wall.
Aperture single with subcircular outline. The embryo is
composed of a spherical proloculus with a diameter of
62µm. The nepionic stage is composed of 9-10 chambers
biserially arranged (Fig. 17W-Z), ephebic of around 3-4
chambers arranged uniserially (Fig. 17Y
-Z). Chamber
lumen not subdivided. Sutures slightly depressed (Fig.
17Y
-Z). Dimorphism not observed.
Remarks. The wall texture and the uniserial ephebic
chamber arrangement suggest that this species belongs to
the subfamily Praerhapydionininae; however, the biserial
nepionic chamber arrangement is different from the
planispiral coiling of the other genera ascribed to this
subfamily. The Cretaceous genus Pseudorhapydionina
de castro, 1971 with nepionic biserial arrangement, has
cribrate aperture and a well-developed exoskeleton. The
lack of sufficient material leads to leave these specimens
undetermined.
Distribution. This species is present in the depositional
sequences Da-2 DS from Urbasa Pass section (Fig. 11),
Se/Th-1 DS from Andia section (Fig. 9) and in Lizarraga
section (Fig. 10) with the species assemblage indicated in
the aforementioned figures.
Biozone. This species was found associated to
assemblages of larger foraminifera that belong to the SBZ
2-SBZ 3 (Fig. 9).

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

Paleocene LBF from the Pyrenean Basin

Family Meandropsinidae henson
Genus Hottingerina drobne
Type species: Hottingerina lukasi, drobne
Hottingerina lukasi
Fig. 16R-U.

drobne,

1975

1975 Hottingerina lukasi. Drobne, p. 224-248; pl. 1;
pl. 2, fig. 1-2; pl. 3; pl. 4, fig. 1-9; pl. 6, fig. 15; pl. 7, fig. 1;
pl. 8; pl. 9; pl. 10, fig. 1-2
2010 Hottingerina lukasi Drobne. Di Carlo et al., p.
64; pl. 6, figs. 1-3.
Material. Samples K 1 and K 2 from Korres section
(Fig. 13).
Description. The test is porcelaneous with lenticular
morphology and planispiral involute growth. The maximum
equatorial diameter observed is about 1.65mm and the wall
is 0.8mm-thick. The endoskeleton consists only of a thin
basal layer. The exoskeleton is composed of short beams
(Fig. 16R-S). The aperture is located at the base of the
septum.
Remarks. The embryonic architecture of this species
could not be observed in the material studied due to the
lack of a centred section.
Distribution. This species has been identified in the
Th-2 DS from the Korres section (Fig. 13) with the species
assemblage indicated in the aforementioned figures.
Biozone. Drobne (1975) and Drobne in Serra-Kiel et
al. (1998) considered this species as belonging to the SBZ
4, from which it can be considered as a marker.
Superfamily Alveolinoidea ehrenberg
Family Alveolinidae ehrenberg
Genus Glomalveolina hottinger
Type species: Alveolina dachelensis schwager
Glomalveolina primaeva (reichel, 1937)
Fig. 18A-H
1937 Alveolina primaeva. Reichel, p. 88-91, fig. 17;
text fig. 15; pl. 9, fig.4-5
1960 Alveolina (Glomalveolina) primaeva reichel.
Hottinger, p. 53-54, text fig. 29 nº 12-14; pl. 1, fig. 3-7
1974 Alveolina (Glomalveolina) primaeva reichel.
Hottinger, p. 34-35; pl. 31, figs. 8-9
1977 Alveolina (Glomalveolina) primaeva reichel.
Drobne, p. 13-14, text fig. 1a-d
1983 Alveolina (Glomalveolina) primaeva reichel.
Rahaghi, p. 49, pl. 21, fig. 1-3
1998 Alveolina (Glomalveolina) primaeva reichel.

27

Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

Sirel, p. 63-64; pl. 29, fig. 5-6, 10-11, 13
2008 Glomalveolina primaeva (reichel). Sirel and
Acar, p. 5; pl. 1, figs. 1-6
2008 Glomalveolina primaeva (reichel). Pignatti et
al. pl. 5, fig. 5; pl. 7, fig. 8a-b
2009 Glomalveolina primaeva (reichel). Sirel, pl. 6,
fig. 12
2010 Glomalveolina primaeva (reichel). Di Carlo et
al. p. 57-58; pl. 5, figs. 1-5, 7-13, 14B, 15, 16
2015 Glomalveolina primaeva (reichel). Sirel, pl. 17,
figs. 5-6, 10-13
2018 Glomalveolina primaeva (reichel). Sirel, pl. 30,
figs. 4-5.
Material. Sample Se 1 from Serraduy section (Fig.
4); samples Cm 3 and Cm 8 from Campo section (Fig. 5);
samples Te 3, Te 4 and Te 5 from Tena section (Fig. 6);
samples An 4, An 6, An 8, An 9 and An 10 from Andia
section (Fig. 9); sample L 25 from Lizarraga section (Fig.
10); samples Le 7 and Le 8 from Leortza section (Fig. 12);
samples Urb 30 and Urb 31 from Urbasa Pass section (Fig.
11) and samples Aix 19, Aix 20, Aix 21, Er 1 and Er 2 from
Aixola-Ermua section (Fig. 14).
Description. The test is porcelaneous with
subspherical morphology. Megalospheric forms show
an embryo formed of a proloculus with a diameter of
about 85-90µm (Fig. 18F G) and a f lexostyle, followed
,
by the nepionic stage consisting of chambers with
streptospiral growth. This nepiont stage has a diameter
of around 220-250µm. The possible microspheric
forms (Fig. 18C, H) show a nepionic stage with a
diameter of about 150µm. In both generations, the
ephebic stage consists of chambers planispirally
arranged subdivided into chamberlets by septula,
which alternate in successive chambers. The maximum
equatorial diameter is around 1.6mm for 7 whorls. The
elongation index is about 1.
Distribution. This species is present in Se/Th-1
DS of the Serraduy section (Fig. 4), Campo section
(Fig. 5), Tena section (Fig. 6), Andia section (Fig. 9),
Lizarraga section (Fig. 10), Urbasa Pass section (Fig.
11), Leortza section (Fig. 12) and Aixola-Ermua section
(Fig. 14) with the species assemblage indicated in the
aforementioned figures. In this latter section samples Er
1 and Er 2 were found in clasts of the calcareous breccia
located in TH-2 DS.
Biozone. According to Hottinger (1960) and Serra-Kiel
et al. (1998) this species is a biostratigraphic marker of the
SBZ 3.
Glomalveolina levis
Fig. 18.9-18.12

hottinger,

1960

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

1960 Alveolina (Glomalveolina) levis. Hottinger, p.
55-57, text fig. 29 nº28; pl. 11-14
1974 Alveolina (Glomalveolina) levis Hottinger, p.
35; Pl. 31, figs. 1-4
Material. Sample Se 2 from Serraduy section (Fig. 4);
sample Cm 10 from Campo section (Fig. 5); sample Te 6
from Tena section (Fig. 6) and samples K 1 and K 2 from
Korres section (Fig. 13).
Description. The test is porcelaneous with ovoidfusiform morphology. The megalospheric form shows an
embryonic stage composed of a proloculus with a diameter
of about 85µm followed by a flexostyle, the nepionic stage
of 2-3 streptospiral whorls and a diameter of about 150µm
(Fig. 18I). The possible microspheric forms (Fig. 18K)
differ from the megalospheric form in possessing a tighter
and smaller nepionic stage. In both generations, the ephebic
stage consists of 7-8 whorls with planispiral chamber
arrangement. The chamber lumen of the ephebic stage is
subdivided into chamberlets by septula. Septula alternate
in position in successive chambers. The elongation index
is 1.3-1.7.
Distribution. This species is identified in Th-2 DS in
the Serraduy section (Fig. 4), Campo section (Fig. 5), Tena
section and in Korres section (Fig. 13) with the species
assemblage indicated in the aforementioned figures.
Biozone. According to Hottinger (1960) and SerraKiel et al. (1998) this species is a biostratigraphic marker
of the upper Thanetian or SBZ 4.
Genus Alveolina d’orbigny
Type species: Oryzaria boscii defrance, in bronn
Alveolina korresensis n. sp. Serra-Kiel and Vicedo
Fig. 18M-Q
Derivation of name. The specific name refers to the
village of Korres.
Holotype. specimen illustrated in Figure 18M. Housed
in the palaeontological collections of the Museu de
Ciències Naturals de Barcelona, accession number MGB
89844 LP01.01.
Paratype. specimen illustrated in Figure 18O. Housed
in the palaeontological collections of the Museu de
Ciències Naturals de Barcelona, accession number MGB
89844 LP01.02.
Type locality. Located in the Korres section, near
the village of Korres. Sample K 2. Coordinates: N 42º 51’
54.54’’/W 2º 26’ 13.62’’
.

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Paleocene LBF from the Pyrenean Basin

FIGURE 18. Paleocene Larger Foraminifera from the Pyrenean Basin. A-H: Glomalveolina primaeva (reichel, 1937) A and D megalospheric forms,
uncentered axial sections; B polyvalent specimens; F-G centered axial sections; C uncentered possible microspheric form; E and H centered
possible microspheric form. Specimens A, B, C and E from sample Cm 8; D, F and G from sample Le 8 and H from sample Urb 31. I-L:
Glomalveolina levis hottinger, 1960 9 centered axial section, megalospheric form; J, K and L megalospheric forms, uncentered axial sections.
All specimens from sample K 2. M-Q: Alveolina korresensis n. sp. centered axial sections of megalospheric forms M Holotype, O Paratype. All
specimens from sample K 3.

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J. Serra-Kiel et al.

Type level. “Assilina beds” upper Thanetian or SBZ 4.
,
Material. Sample K 2 from the Korres section (Fig.
18M).
Diagnosis. Porcelaneous test, axially elongated
morphology with rounded poles. Multiple apertures.
Microspheric forms not found. Megalospheric form with
a nepionic stage that consists of a proloculus of circular
outline in axial section and a diameter of 140-190μm
followed by a flexostyle. Nepionic stage composed of 2
tight whorls, ephebic stage formed by 5 whorls. Maximum
axial and equatorial diameters observed of 1.3-1.6mm and
0.5-07mm respectively, with an elongation index of 2.0-2.1.
Chamberlets in the nepionic stage showing a subrectangular
outline in axial section becoming oval to subrectangular in
ephebic whorls. Endoskeleton consisting of a thin basal
layer and septula subdividing chambers into chamberlets.
Septula alternating position in successive chambers.
Remarks. This species is the oldest representative of
the genus Alveolina in the Paleocene from the Pyrenean
basin. It differs from A. vredenburgi da
vies and pinfold,
1937 (=ex A. cucumiformis) in the smaller size of the test
and in the thickness of the basal layer.
Distribution. This species is present in Th-2 DS in
the Korres section (Fig. 13) with the species assemblage
indicated in the aforementioned figure.
Biozone. According to the foraminiferal assemblage, A.
korresensis can be considered as belonging to the SBZ 4.
Class GLOBOTHALAMEA pawlowski et al.,
Order “TEXTULARIIDA” (delage and hérouard
partim
Superfamily Eggerelloidea cushman
Family Valvulinidaea berthelin
Subfamily Valvulininaea berthelin
Genus Goesella cushman
Type species: Clavulina rotundata cushman
Goesella sp.
Fig. 19A-E
2001 Goesella sp. Ogorelec et al., pl. 10, fig. 6
Material. Sample Urb 20 from Urbasa Pass section
(Fig. 11); sample Abau 1 from Abaurrea section (Robador,
2005, Annex, p. 50-51) and sample Bin 3 from Foz de
Biniés section (Robador, 2005, Annex, p. 90-91).
Description. The specimens show a finely agglutinated
test of small size. The embryo consists of an ovoidal
proloculus with a maximum diameter of around 135-120μm

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and minimum of around 80-110μm. The neanic stage is
composed of biserial chambers followed by the ephebic
stage characterized by chambers arranged uniserially.
Distribution. This species has been identified in the
DS Da-2 of the Urbasa Pass section (Fig. 11) and in samples
from the lower part of the sections of Abaurrea and Foz de
Biniés (see Robador, 2005). It is associated with species
without biostratigraphical interest such as Lenticulina sp.
and Robulus sp.
Biozone. Ogorelec et al. (2001) consider that this
species belongs to the SBZ 2. This zonal assignment is
in accordance with the data obtained in the Urbasa Pass
section (Fig. 11). Its occurrence in the lower levels of the
sections of Abaurrea and Foz de Biniés studied by Robador
(2005) indicates that this species could also be present in
SBZ 1.
Superfamily Coscinophragmatoidea thwlmann
Family Haddoniidae chapman
GENUS Haddonia chapman
Type species: Haddonia torresiensis, chapman
Haddonia praeheissigi
1977
Fig. 19F

samuel, kohler and borza,

1972 Haddonia sp. Samuel et al., pl. 2, figs. 1-2; pl. 3,
figs.1-2
1977 Haddonia praeheissigi n. sp., Samuel et al. pl.
48, figs. 1, 2; pl. 49, figs.1, 2; pl. 50. figs. 1, 2
1988 Haddonia sp., Drobne et al., pl. 26, fig. 5
2008 Haddonia praeheissigi samuel, köhler and
borza. Pignatti et al., pl. 7, fig. 5
2010 Haddonia praeheissigi samuel, köhler and
borza. Di Carlo et al., p. 50-51; pl. 1, fig. 1; pl. 3, figs.
1-2
Material. Samples L 8 and L 9 from Lizarraga section
(Fig. 10); samples Urb 2, Urb 4, Urb 6, Urb 7, Urb 13, Urb
25, Urb 27 and Urb 28 from Urbasa Pass section (Fig. 11);
sample Le 2 from Leortza section (Fig. 12) and samples
Aix 5 and Er 5 from Aixola-Ermua section (Fig. 14).
Description. The wall is composed of two layers, the
external layer consists of coarsely agglutinated particles,
mainly of quartz, calcareous cement, and a thin inner
layer formed of micrite, which is poorly preserved in the
material studied. The morphology is irregular, showing
protuberances on the external surface. Chambers are also
irregular in shape.
Distribution and age. The morphology of this species
suggests that it lived attached to the substrate, probably

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Paleocene LBF from the Pyrenean Basin

FIGURE 19. Paleocene Larger Foraminifera from the Pyrenean Basin. A-E: Goesella sp. longitudinal sections. Specimens A, B and E from sample
Abau 1; C from sample Bin 3 and D from sample Urb 20. F: Haddonia praeheissigi, samuel et al., 1977 specimen from sample Er 5. G: Coscinophragma
cribosum (reuss, 1846) specimen from sample Lech 12. H: Kolchidina cf. paleocenica (cushman, 1947) uncentered longitudinal section, specimen
from sample Bur 3. I: Placopsilina cenomana d’orbigny, 1850 specimen from sample Le 2. J: Popovia cf. plana (bykova, 1939) equatorial section,
specimen from sample Orio 3. K-N: Vania anatolica sirel and gündüz, 1985. K-M uncentered axial sections; N equatorial section; Specimens K and
M from sample Cm 5; L from sample Cm 7 and N from sample Le 8. O-T: Coskinon rajkae (hottinger and drobne, 1980) O oblique-basal section; P
section parallel and close to axis; Q oblique section close to axial plane; R-T axial sections. Specimen O from sample Te 4; P from sample Cm 5; Q
from sample Cm 8; R and S from sample Le 8 and T from sample Bur 7. U-C’: Fallotella alavensis mangin, 1954. U, V and A’ longitudinal sections
parallel to axial plane; W and Y oblique basal sections; X and B’ basal sections; Z oblique section; C’ longitudinal oblique section. Specimens U and
A’ from sample Le 5; V from sample Cm 8; W, Y and Z from sample Le 8; X and B’ from sample Le 7 and C’ from sample Bur 7. D’-I’: Cribrobulimina
carniolica hottinger and drobne, 1980. D’, H’ and I’ no centered sections parallel to axis; E’ basal section showing the foramina in the septum; F’ and
G’ oblique sections. Specimens D’, and G’ from sample Cm 5; E’, F’ and H’ from sample Cm 8 and I’ from sample Coll 3.

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to “seagrasses” and substrates from reef environments
developed in the inner shelf. It has no biostratigraphic
significance.
Family: Coscinophragmatidae thalmann
GENUS Coscinophragma thalmann
Type species: Lichenopora cribosa, reuss
Coscinophragma cribosum (reuss, 1846)
Fig. 19G
1972 Coscinophragma cribosum (reuss, 1846).
Samuel et al., pl. 4, fig. 1-2; pl. 5, fig. 1
Material. Sample Lech 12 from Lecherín section
(Robador, 2005, Annex, p. 24-25).
Description. The shell is agglutinated, flattenedirregular in shape, with branches. The ephebic stage shows
chambers in uniserial distribution. The wall is composed
of three layers: the outer part consisting of a homogeneous
calcareous layer, the inner layer composed of large quartz
grains and the median layer is pierced by canaliculi
perpendicular to the surface of the test. The apertures are
cribrate.
Distribution and age. The morphology of this
species suggests that it lived attached to the substrate,
probably on the “seagrass” and on otherv substrates of reef
environments of the inner shelf. It has no biostratigraphic
significance.
Superfamily: Lituoloidea de blainville
Family: Lituolidae de blainville
Subfamily: Lituolinae de blainville
GENUS Kolchidina, morozova
Type species: Ammobaculites manyschensis, bykova
Kolchidina cf. paleocenica (cushman, 1947)
Fig. 19H
1998 Kolchidina paleocenica (cushman). Sirel, p. 35365; pl. 1, fig. 19-24
2009 Kolchidina paleocenica (cushman). Sirel, pl. 2,
figs. 15-16
2015 Kolchidina paleocenica (cushman). Sirel, pl. 4,
figs. 1-6
2018 Kolchidina paleocenica (cushman). Sirel, pl. 48,
figs. 1-6
Material. Sample Bur 3 collected by Robador from
Burgi section (Robador, 2005, Annex, pp. 78-79).
Description. The test is coarsely agglutinated
with a thick wall. The nepionic stage consists of

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a proloculus followed by planispirally arranged
chambers. The ephebic stage, consisting of uniserially
arranged chambers, is not present in the single
specimens found.
Remarks. The specific determination of this taxon
has been left open since the ephebic stage could not be
observed.
Distribution. In the material studied K. cf. paleocenica
is associated with M. minutus and T. madrugaensis.
Biozone. According to Sirel (1998) this species occurs
in the Danian from Turkey. These data and the results
presented here indicate that its biostratigraphic range is
within SBZ 3.
Family: Placopsilinidae rhumbler
Subfamily: Placopsilininae rhumbler
GENUS Placopsilina D’orbigny
Type species: Placopsilina cenomana, D’orbigny
Placopsilina cenomana D’orbigny, 1850
Fig. 19I
1972 Placopsilina cenomana D’orbigny. Samuel et
al., pl. 1, fig. 1-2
Material. Samples Urb 2 and Urb 3 from Urbasa Pass
section (Fig. 11) and sample Le 2 from Leortza section
(Fig. 12).
Description. The test is flattened with irregular
morphology. Wall thick and formed of coarse agglutinated
particles. Each chamber overlaps the previous, forming a
relief on the surface of the test. The apertures are located
in the base of the septum and with a slight lip. The septal
sutures are well-marked.
Distribution.
The morphology of this species
suggests that this species lived attached to the substrate,
probably on the “seagrass” and on other substrates from
reef environments of the inner shelf. This species is found
in Ma-Da DS from Urbasa Pass section and in Se/Th-1 DS
from Leortza section.
Biozone. According to the data presented herein, the
biostratigraphic range of this species is from SBZ 1 to
SBZ 3.
Superfamily: Loftusioidea brady
Family: Cyclamminidae marie
Subfamily: Alveolophragmiinae saidova
GENUS Popovia, suleymanov
Type species: Alveolophragmium planum, bykova

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J. Serra-Kiel et al.

Popovia cf. plana (bykova, 1939)
Fig.19J
1998 Popovia plana (bykova). Sirel, p. 37-38; pl. 35,
figs. 37-38
2004 Popovia sp., Sirel, pl. 9, figs. 18-19
2015 Popovia plana (bykova). Sirel, pl. 5, figs. 13-14
Material. Sample Orio 3 from a pebble taken from
Orio section (p. 248, fig. 75, Baceta, 1996).
Description. The wall is agglutinated, thick with
an alveolar texture. The apertures in nepionic stage are
interiomarginal in position. The ephebic chambers have a
uniserial arrangement.
Remarks. The specific identification of this species
has been left in open nomenclature since the nepionic stage
could not been observed.
Distribution. This species has been found associated
with M. minutus and T. madrugaensis.
Biozone. According to Sirel (1998) this species occurs
in the Thanetian from Turkey associated with G. primaeva,
F. kochanskae and I. sinjarica. These data and the results
presented here indicate that its biostratigraphic range is
SBZ 3.
Family: Spirocyclinidae munier-chalmas
GENUS Vania sirel and gündüz
Type species: Vania anatolica sirel and gündüz
Vania anatolica sirel and gündüz, 1985
Fig. 19K-N
1971 Broeckinella arabica henson. Drobne and
Hottinger, p. 9, text fig. 2d, e; 3a-d; pl. 3, 4
1978 Broeckinella arabica henson. Rahaghi, p. 39;
pl. 1, fig. 1, 2
1985 Vania anatolica Sirel and Gündüz, p. 20-24; text
fig. 2A, B; pl. 1 fig. 1-3, pl. 2, fig. 1-6, pl. 3, fig. 1-7
1989 Broeckinella arabica henson. Kuss and Leppig,
p. 312, fig. 5k
1998 Vania anatolica sirel and gündüz. Sirel, p. 4344, pl. 6, fig. 9-14
2004 Vania anatolica sirel and gündüz. Sirel, p. 2224; pl. 24, figs. 1-3; pl. 25, figs. 1-6; pl. figs. 1-7; textfig. 10 A, B
2008 Vania anatolica sirel and gündüz. Pignatti et
al., pl. 7, figs. 3-4
2010 Vania anatolica sirel and gündüz. Di Carlo et
al., p. 51; pl. 3, figs. 3-7ª, 8
2012 Vania anatolica sirel and gündüz. Sirel, pl. 8,
figs. 10-15

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2015 Vania anatolica sirel and gündüz. Sirel, pl. 27,
figs. 10-15 and pl. 50, figs. 10-15
Material. Samples Cm 3, Cm 5 and Cm 7 from Campo
section (Fig. 5); sample Te 4 from Tena section (Fig. 6);
sample G 3 from Garralda section (Fig. 7); sample An 10
from Andia section (Fig. 9) and samples Le 3, Le 6, Le 7
and Le 8 from Leortza section (Fig. 12).
Description. The test is agglutinated with discoidal
morphology. In the material studied, the embryonic and
nepionic stages could not be observed. The chambers of
the ephebic stage are annular. The exoskeleton consists of
beams and rafters, forming a polygonal network. Foramina
are arranged in rows with radial axes in successive chambers.
Distribution. This species is found in the Se/Th-1
DS of the Campo section (Fig. 5), Tena section (Fig. 6),
Garralda section (Fig. 7), Andia section (Fig. 9) and Leortza
section (Fig. 12) with the species assemblage indicated in
the aforementioned figures.
Biozone. According to Sirel (1998) this species
occurs in the Thanetian associated with G. primaeva,
G. dachelensis, M. juliettae, C. rajkae and I. sinjarica.
Later, Serra-Kiel et al. (1998) considered this species as
belonging to SBZ 3, from which it can be considered a
marker.
Superfamily Ataxophragmioidea schwager
Family Coskinolinidae moullade
GENUS Coskinon hottinger and drobne
Type species: Coskinolina (Coskinolina) rajkae,
hottinger and drobne

Coskinon rajkae (hottinger and drobne, 1980)
Fig. 19O-T
1980 Coskinolina (Coskinon) rajkae. Hottinger and
Drobne, p. 45-46, text fig. 2; pl. 2, fig. 2-4; pl. 12. figs.
1-28
1988 Coskinon rajkae (hottinger and drobne).
Drobne et al., pl. 26, fig. 7, 8
1998 Coskinolina (Coskinon) rajkae (hottinger and
drobne). Sirel, p. 47, pl. 9, fig. 6-10
2001 Coskinon rajkae (hottinger and drobne).
Ogorelec et al., pl. 10, fig. 7
2001 Coskinon rajkae (hottinger and drobne).
Özgen and Akyazi, pl. 1, fig. 9-11
2008 Coskinon rajkae (hottinger and drobne).
Pignatti et al., pl. 7, figs. 6, 9b
2010 Coskinon rajkae (hottinger and drobne). Di
Carlo et al., p. 51-52; pl. 3, figs. 10, 11, 13; pl. 5, fig.
25A
2013 Coskinon rajkae (hottinger and drobne).

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Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

Sirel, p. 34, pl. 5, figs. 1-5
2015 Coskinon rajkae (hottinger
Sirel, p. 28, figs. 1-5
2018 Coskinon rajkae (hottinger
Sirel, p. 56, figs. 1-5

and drobne).
and drobne).

Material. Samples Cm 5, Cm 7 and Cm 8 from Campo
section (Fig. 5); samples Te 4 and Te 5 from Tena section
(Fig. 6); sample An 6 from Andia section (Fig. 9); sample
L 26 from Lizarraga Section (Fig. 10); samples Le 5, Le 7
and Le 8 from Leortza section (Fig. 12) and sample Bur 7
from Burgi section (Robador, 2005, Annex, p. 76-77).
Description. The test is agglutinated with high-conical
morphology and cribrate aperture. The wall is thick and
displays a pseudokeriothecal texture. The embryo could not
be adequately observed. It is followed by chambers with
trochospiral arrangement. The ephebic stage is composed
of 8-10 chambers arranged uniserially. The apertural face
is convex in early growth stages, becoming slightly convex
or flat in the ephebic stage. The endoskeleton is composed
of pillars arranged discontinuously from one chamber to
the next. At a cone diameter of 0.9mm, the axial length is
1.2mm. The axial plane cuts 5-6 pillars in adult chambers.
Chamber sutures are slightly depressed.
Distribution. This species is present in the Se/Th-1 DS
of the Campo section (Fig. 5), Tena section (Fig. 6), Andia
section (Fig. 9), Lizarraga section (Fig. 10) and in Leortza
section (Fig. 12), with the species assemblage indicated in
the aforementioned figures.
Biozone. Hottinger and Drobne (1980), Drobne et al.,
(1988), Sirel (1998), Ogorelec et al. (2001) and Özgen
and Akyazi (2001) reported this species as belonging to
the Glomalveolina primaeva Zone. This species can be
considered as a marker of the SBZ 3.
Superfamily Orbitolinoidea martin
Family Orbitolinidae martin
Subfamily Dictyoconinae moullade
GENUS Fallotella mangin
Type species: Fallotella alavensis mangin
Fallotella alavensis mangin, 1954
Fig. 19U-C’
1954 Fallotella alavensis sp., Mangin, p. 209; text
figs. 1-3; pl. 3, figs. 1-6
1980 Fallotella (Fallotella) alavensis Mangin.
Hottinger and Drobne, p. 50-51, pl. 14, fig. 1-27
1983 Fallotella alavensis Mangin. Rahaghi, p. 28-29,
pl. 2, fig. 1-5
2001 Fallotella alavensis Mangin. Özgen and Akyazi,
pl. 1, fig. 4-6

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2008 Fallotella alavensis Mangin. Pignatti et al., pl.
5, fig. 2; pl. 7, figs. 2, 7a
2010 Fallotella alavensis Mangin. Di Carlo et al., p.
52; pl. 3, figs. 14-22
Material. Samples Cm 3 and Cm 8 from Campo
section (Fig. 5); samples Te 3, Te 4 and Te 5 from Tena
section (Fig. 6); samples Le 5, Le 7 and Le 8 from Leortza
section (Fig. 12); samples Aix 19 and Aix 20 from AixolaErmua section (Fig. 14) and sample Bur 7 from Burgi
section (Robador, 2005, Annex, p. 76-77).
Description. The test is agglutinated with high-conical
morphology. The cone surface is smooth. The wall is thin
without ultrastructure. The maximum cone-length observed
is around 1.8mm with 15-16 uniserial chambers. The
maximum diameter measured of an adult chamber is around
1.7mm. The embryonic stage has not been adequately
observed. The nepionic stage is composed of chambers
in trochospiral arrangement. The neanic stage consists of
chambers arranged uniserially. There are 10-12 chambers
per millimetre of axial length. The exoskeleton appears first
in the neanic stage and consists of beams. The endoskeleton
is composed of pillars with a subcircular horizontal section,
except below the ceiling of the chamber where pillars have
an elliptical shape in transverse section. Pillars alternate in
position from one chamber to the next. The apertural face is
slightly convex, and the aperture is cribrate.
Distribution. This species is present in the Se/Th-1
DS of the Campo section (Fig. 5), Tena section (Fig. 6),
Leortza section (Fig. 12) and in Aixola-Ermua section
(Fig. 14) with the species assemblage indicated in the
aforementioned figures.
Biozone. This species was described by Mangin (1954)
in the Paleocene from the Pyrenean basin. Later, Hottinger
and Drobne (1980) and Serra-Kiel in Robador et al. (1991a,
b) found it associated with G. primaeva, C. rajkae and V.
anatolica, pointing it as a marker of the SBZ 3.
Superfamily: Eggerelloidea cushman
Family: Valvulinidae berthelin
Subfamily: Valvulininae berthelin
GENUS Cribrobulimina cushman
Type species: Valvulina mixta parker and jones
Cribrobulimina carniolica hottinger and drobne,
1980
Fig. 19D’-I’
1980 Cribrobulimina carniolica, Hottinger and
Drobne, p. 219-221; pl. 3, fig. 1-11; pl. 2, fig. 1, 3
1988 Cribrobulimina carniolica hottinger and
drobne. Drobne et al., pl. 26, fig. 10-11

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J. Serra-Kiel et al.

2001 Cribrobulimina carniolica hottinger and
drobne. Özgen and Akyazi, pl. 1, fig. 7, 8
2010 Cribrobulimina aff. carniolica hottinger and
drobne, 1980. Di Carlo et al., p. 53; pl. 3, figs. 9, 12
Material. Sample Se 1 from Serraduy section (Fig.
4); samples Cm 5 and Cm 8 from Campo section (Fig. 5);
samples Te 3, Te 4 and Te 5 from Tena section (Fig. 6);
samples Le 6, Le 7 and Le 8 from Leortza section (Fig. 12)
and sample Coll 3 from Collarada section (Robador, 2005,
Annex, p. 22-23).
Description. The test is agglutinated with low-conical
morphology. The wall is thick with pseudokeriothekal
texture. Chambers are inflated and arranged trochospirally.
The sutures are strongly depressed. The apertures are
interiomarginal in position in the early stage of growth,
later becoming cribrate with multiple large apertures.
Distribution. This species is present in Se/Th-1 DS
in the Serraduy section (Fig. 4), Campo section (Fig. 5),
Tena section (Fig. 6) and Leortza section (Fig. 12) with the
species assemblage indicated in the aforementioned figures.
Biozone. Hottinger and Drobne (1980) identified this
species in Thanetian rocks associated to G. primaeva, F.
alavensis, C. rajkae and V. anatolica (=ex Broeckinella
arabica). Later, Serra-Kiel et al., (1998) considered it as a
marker of the SBZ 3.
Order: Rotaliida (delage and hérouard)
Superfamily: Rotalioidea ehrenberg
Family: Elphidiidae galloway
GENUS Bangiana drobne, ogorelec and
riccamboni

Type species: Bangiana hanseni drobne, ogorelec
and riccamboni

Bangiana
riccamboni,

hanseni
2007

drobne,

ogorelec

Distribution. This species is present in the Da-1 DS
from Tena section (Fig. 6). Da-2 DS from Urbasa Pass
section (Fig. 11) associated with H. paleocenica, H.
elongata and Goesella sp. and overlaying M. globularis
and O. moorkensii. Finally, this species is also present in
DA-1 DS and DA-2 DS from Urrobi section (Fig. 8).
Biozone. Drobne et al. (1988), Serra-Kiel et al.
(1998) and Drobne et al. (2007) considered this species as
a marker of the SBZ 1. However, in the material studied it
is associated with foraminifera of SBZ 2. Consequently, its
range should be extended to this latter biozone.
Family: Rotaliidae ehrenberg
Subfamily: Rotaliinae ehrenberg
GENUS Rotorbinella bandy
Type species: Rotorbinella colliculus bandy
Rotorbinella hensoni (smout, 1954)
Fig. 20P-S
1954 Rotalia hensoni. Smout, p. 45; pl. 15, fig. 8
1973 Discorbis perovalis (terquem). Ferrer et al., p.
38; pl. 37, figs. 1, 3
2006 Rotorbinella sp. hottinger, p. 86; pl. 2, figs. 1116
2014 Rotorbinella hensoni (smout, 1954). Hottinger,
p. 24; Figs. 3.2, 3.3A-F; pl. 3.2, figs. 1-13
2014 Rotorbinella detrecta. Hottinger, p. 26, pl. 3.4AH
Material. Sample An 8 from Andia section (Fig. 9);
samples Urb 5, Urb 6 and Urb 9 from Urbasa Pass section
(Fig. 11) and samples Ur 8, Ur 10, Ur 11, Ur 16 and Ur 17
from Urko section (Fig. 15).

and

Fig. 20A-O
1988 Protelphidium sp. Drobne et al., pl. 25, figs. 8-11
2001 Bangiana hanseni. Ogorolec et al., pl. 9, figs.
16-20
2007 Bangiana hanseni. Drobne et al., p. 16-24; pls.
1-12
Material. Sample Te 1 from Tena section (Fig. 6); samples
Urrob 1 and Urrob 2 from Urrobi section (Fig. 8) and samples
Urb 12, Urb 16 and Urb 21 from Urbasa Pass section (Fig. 11).
Description. The lamellar-perforated test is small,
biumbilical, depressed with a planispiral involute chamber
arrangement. The wall is thin. The chambers are inflated

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increasing in size through ontogeny. The apertures are
located at the base of the apertural face.

Description. Small size, bilamellar-perforate test. The
morphology is low-trochospiral with a rounded periphery.
The ventral and dorsal sides have high convexity. The
umbo is pronounced with a massive plug. The diameter
of the proloculus is small (about 20µm). The diameter of
the shell with 2 whorls is about 0.58mm and the height is
0.35mm.
Distribution. This species is present in the Da-1 DS
from Urbasa Pass section (Fig. 10), and Urko section (Fig.
15), Da-2 DS from Urko section (Fig. 16), Se/Th-1 DS
from Andia section (Fig. 9) with the species assemblage
indicated in the aforementioned figures.
Biozone. According to Hottinger (2014) and Vicedo
et al. (2019) its biostratigraphic range is SBZ 2 to SBZ 3.

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Paleocene LBF from the Pyrenean Basin

FIGURE 20. Paleocene Larger Foraminifera from the Pyrenean Basin. A-O: Bangiana hanseni drobne et al., 2007. A-B, E-G, I-O and Q axial sections.
C-D oblique sections; H-P equatorial sections. Specimens A, B, E and K from sample Urrob 1; C, D, F-I and L-O from sample Urb 12 and J from
sample Te 1. P-S: Rotorbinella hensoni (smout, 1954) axial sections. Specimen P from sample Ur 8; Q from sample Ur 10; R and S from sample
An 8. T-V: Rotorbinella skourensis (pfender, 1938) axial sections. Specimens T, U from sample Ur 8 and V from sample Ur 10. W-X: Pyrenerotalia
depressa hottinger, 2014 slightly oblique axial sections. Specimens from sample Ur 8. Y-A’: Rotospirella conica (smout, 1954). Y-Z adaxial sections.
A’ oblique section. Specimen Y from sample Salv 4 and Z and A’ from sample L 3. B’-H’: Redmondina henningtoni hasson, 1985. B’-C’ no centered
sections perpendicular to coiling axis. D’-F’ adaxial sections. G’ oblique section to coiling axis, Specimen B’ from sample K 2 and C’-H’ from
sample Cm 8. I’-P’: Kathina aquitanica hottinger, 2014. I’-L’ and O’ centred axial section; M’ and P’ uncentered axial sections and N’ oblique
section. Specimens I’, J’, M’, N’ from sample An 10; K’ from sample An 9; O’ from sample Aix 19; P’ from sample L 24 and L’ from sample Cm 3.
Q’-X’: Kathina pernavuti sirel, 1972 axial sections. Specimen Q’ from sample Salv 5 and R’-X’ from sample Cm 8. Y’-B’’: Elazigina dienii (hottinger,
2014) all specimens in axial sections. Specimens Y’-A’’ from sample Aix 9 and B’’ from sample L 3. Abbreviations: pr: proloculus, um: umbilical
umbo, up: umbilical plate, ilsp: intraseptal interlocular space, spc: spiral canal, fu: funnel, fea: feathering interlocular space, fol: folium, pil: pile,
fol: folium, spsut: spiral suture.

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association of P. depressa with O. moorkensii implies a
biostratigraphic range equivalent to the SBZ 2.

Rotorbinella skourensis (pfender, 1938)
Fig. 20T
-V
2009 Rotalia trochidiformis lamarck. Afzal et al.,
pl. 1, fig. 12
2014 Rotorbinella skourensis (pfender, 1938).
Hottinger, p. 24; pl. 3.1, figs. 1-18
Material. Sample Cm 3 from Campo section (Fig. 5);
sample L 26 from Lizarraga section (Fig. 10) and samples
Ur 8 and Ur 10 from Urko section (Fig. 15).
Description. Bilamellar-perforate test. The morphology
is low-trochospiral with rounded periphery. The ventral side
has high convexity. The umbo is pronounced. The diameter
of the proloculus is around 60µm. The adult shell has a
diameter of around 0.50mm with 3 whorls and a height of
0.33mm.
Distribution. This species is present in the Da-1 DS
from Urko section (Fig. 15), Se/Th-1 DS from Campo
section (Fig. 5) and in Lizarraga section (Fig. 10) with
the species assemblage indicated in the aforementioned
figures.
Biozone. According to Hottinger (2014) R. skourensis
belongs to the SBZ 3-SBZ 4. However, the presence of this
species in the Da-1 DS in the Urko section (Fig. 14) implies
that its biostratigraphic range ranges from SBZ 2 to SBZ 4.
GENUS Pyrenerotalia boix, villalonga, caus and
hottinger

Type species: Pyrenerotalia longifolia
villalonga, caus and hottinger
Pyrenerotalia depressa
Fig. 20W-X

hottinger,

Subfamily: Praelockhartiinae vicedo and roblessalcedo

GENUS Rotospirella hottinger
Type species: Lockhartia conica, smout
Rotospirella conica (smout, 1954)
Fig. 20Y ’
-A
1954 Lockhartia conica. smout, p. 53; pl. 4, figs.1-3
1991 Lockhartia conica smout. Wan, p. 162; pl. 1,
figs. 28-29
2014 Rotospirella conica (smout). hottinger, p. 2931; fig. 3.4
Material. Sample L 3 from Lizarraga section (Fig.
10) and sample Salv 4 from Salvatierra de Esca section
(Robador, 2005, Annex, p. 92-95).
Description. Test with coarsely perforated, bilamellar
wall and conical morphology. The chambers are arranged
trochospirally. The dorsal side is smooth without
ornamentation; the chambers sutures flush. The ventral side
is covered by long and delicate folia. Umbilicus composed
of piles and funnels. The diameter of the proloculus is
around 110µm.
Distribution. This species has been identified in the
Da-2 DS of the Lizarraga section (Fig. 10) with the species
assemblage indicated in the aforementioned figure.

boix,

2014

2014 Pyrenerotalia depressa. Hottinger, p. 26, 29-30;
fig. 3.5A-I
Material. Sample Ur 8 from Urko section (Fig. 15).
Description. Test with coarsely perforated wall. The
morphology is lenticular with a trochospiral chamber
arrangement and rounded periphery. The folia overlap the
adaxial part of the umbilicus (Fig. 20). The diameter for 3
whorls is around 1mm and the height 0.45mm.
Distribution. This species has been identified in Urko
section (Fig. 15) in Da-1 DS with the species assemblage
indicated in the aforementioned figure.
Biozone. P. depressa was studied by Hottinger
(2014) without specifying its biostratigraphic range. The

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Biozone. The larger foraminifera associated to
this species suggests a biostratigraphic range within
SBZ 2.
Subfamily: Redmondininae hottinger
GENUS Redmondina hasson
Type species: Redmondina henningtoni hasson
Redmondina henningtoni hasson, 1985
Fig. 20B’-H’
1985 Redmondina henningtoni. Hasson, p. pl. 4.1,
figs. 1-19; pl. 4.2, figs. 1-13
2000 Redmondina henningtoni hasson. Peybernès et
al., p. 46, figs. 67 and 8
2014 Redmondina henningtoni hasson. Hottinger, p.
41; pl. 4.1, figs 1-19; pl. 4.2, figs. 1-13
Material. Samples Cm 4, Cm 5 and Cm 8 from Campo
section (Fig. 5); sample Te 5 from Tena section (Fig. 7);
samples Le 7 and Le 8 from Leortza section (Fig. 12) and
K 2 from Korres section (Fig. 13).

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J. Serra-Kiel et al.

Description. Bilamellar perforate shell with
trochospiral chamber arrangement. The chamber wall
shows coarse pores and rounded periphery. The umbilical
filling is reduced with short and folded folia. The proloculus
diameter is around 50µm. The diameter of the shell varies
from 0.70 to 0.80mm with 3 whorls and the height from
0.40 to 0.50mm.
Distribution. This species is present in the Se/Th-1 DS
from Campo section (Fig. 5); Tena section (Fig. 6) and in
Leortza section (Fig. 12) and in the Th-2 DS from Korres
section (Fig. 13) with the species assemblage indicated in
the aforementioned figures.
Biozone.
According
Hottinger
(2014)
the
biostratigraphic range of this species extends from SBZ 3
to SBZ 6.
Subfamily: Kathininae hottinger
GENUS Kathina smout
Type species: Kathina delseota smout
Kathina aquitanica hottinger, 2014
Fig. 20I’-P’
1998 ?Kathina gr. pernavuti sirel. Accordi et al., p.
200; pl. 16, fig. 2
2006 Kathina sp. Hottinger, p. 87; pl. 2, figs. 17-19
2014 Kathina aquitanica. Hottinger, p. 100; pl. 4.4,
figs. 1-14; pl. 6.2, figs. 1-7

Paleocene LBF from the Pyrenean Basin

(Fig. 9), Lizarraga section (Fig. 10) and Aixola-Ermua
section (Fig. 14).
Biozone. K. aquitanica was characterized by
Hottinger (2014) and attributed to the SBZ 3. The
presence of this species in the Da-2 DS in AixolaErmua section (Fig. 13) and in Urko section (Fig. 15)
implies extending its biostratigraphic range from SBZ
2 to SBZ 4.
Kathina pernavuti sirel, 1972
Fig. 20Q’-X’
1972 Kathina pernavuti. Sirel, p. 289; pl. 5, fig. 7
2014 Kathina pernavuti sirel. Hottinger, p. 101; pl.
6.4, figs. 1-22
Material. Sample Cm 8 from Campo section (Fig.
5) and sample Salv 5 from Salvatierra de Esca section
(Robador, 2005, Annex, p. 92-95).
Description. Bilamellar perforate, small-size test
with low-trochospiral growth. The dorsal side is more
convex than the ventral one (Fig. 20Q’-R’, V’, 20W’).
The solid umbilical structure is pierced by funnels (Fig.
20R’, S’, V’). The folia can be clearly observed (Fig.
20R’, X’). The diameter of the proloculus is around
40-50µm (Fig. 20Q’). The adult shell has a diameter
of around 1mm with 3 whorls and height of around
0.5mm.

Material. Sample Se 1 from Serraduy section (Fig.
4); sample Cm 3 from Campo section (Fig. 5); sample
G 3 from Garralda section (Fig. 7); samples An 3, An 9
and An 10 from Andia section (Fig. 9); sample L 24 from
Lizarraga section (Fig. 10); sample Aix 19 from AixolaErmua section (Fig. 14) and sample Ur 18 Urko section
(Fig. 15).

Distribution. This species has been identified in the
Campo section (Fig. 4) in Se/Th-1 DS with the species
assemblage indicated in the aforementioned figure.

Description. Bilamellar perforate specimens
with trochospiral growth and lenticular morphology.
The ventral side is more convex than the dorsal side
(Fig. 20I’, M’, P’). The test surface is smooth, without
ornamentation. The umbilicus, with a structure typical
of the Kathininae, shows a solid umbilical mass pierced
by funnels (Fig. 20L O’). Specimens show a small
’,
umbilical plate and a tiny spiral canal (Fig. 20O’). The
proloculus has a diameter of 70µm. The diameter of the
adult shell varies from 0.7 to 1mm with 2-3 whorls and
the height from 0.5 to 0.6mm.

GENUS Elazigina sirel
Type species: Kathina subsphaerica, sirel

Distribution. This species occurs in the depositional
sequence Da-2 DS from Urko section (Fig. 15) and in
the Se/Th-1 DS from Serraduy section (Fig. 4), Campo
section (Fig. 5), Garralda section (Fig. 7), Andia section

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Biozone. According to Hottinger (2014) the
biostratigraphic range of this species extends from SBZ 3
to SBZ 4.

Elazigina dienii (hottinger, 2014)
Fig. 20Y’-B’’
2000 “Plumokathina” dienii. Peybernès et al., p. 4647; fig. 5
2001 Plumokathina sp. Ogorolec et al., pl. 10, fig. 8
2014 Plumokathina dienii Hottinger, p. 110; figs. 3.5J,
6.1A-N; pl. 6.8, figs. 1-21
Material. Sample G 2 from Garralda section (Fig. 7);
sample L 3 from Lizarraga section (Fig. 10); sample Aix 9
from Aixola-Ermua section (Fig. 14) and samples Ur 15,
Ur 17 and Ur 18 from Urko section (Fig. 15).

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J. Serra-Kiel et al.

Description. Small, bilamellar perforate specimens
with sharp periphery (Fig. 20Y’-A and low-trochospiral
’’)
growth. The dorsal and ventral sides are convex. The
umbo has funnels (Fig. 20Z’ B’’). The feathering of the
,
intraseptal interlocular space is marked (Fig. 20Y’-B’’).
The proloculus is between 50-120µm in diameter (Fig.
20A The adult shell has a diameter of around 1.0mm
’’).
with 3 whorls and a height of 0.5-0.6mm.
Distribution. This species is present in the Da-2 DS
from Garralda section (Fig. 7), Lizarraga section (Fig.
10), Aixola-Ermua section (Fig. 14) and in Urko section
(Fig. 15) with the species assemblage indicated in the
aforementioned figures.
Biozone. According to Hottinger (2014) this species
belongs to SBZ 2. It can be considered as a marker of this
biozone.
Elazigina lenticula (hottinger, 2014)
Fig. 21A
1972 Rotalia? sp. 3. Samuel et al. pl. 38, figs. 1-2
1998 Plumokathina sp. Accordi et al. pl. 14, fig. 6
2014 Plumokathina lenticula Hottinger, p. 110 and
117; pl. 6.9, figs. 1-6; pl. 6.10, figs. 1-12
Material. Samples An 3, An 8 and An 9 from Andia
section (Fig. 9); sample L 26 from Lizarraga section (Fig.
10); samples K 1 and K 2 from Korres section (Fig. 13) and
samples Er 4 and Er 5 from Aixola-Ermua section (Fig. 14).
Description. The test is bilamellar-perforated with
trochospiral chamber arrangement. The morphology is
lenticular with inflated umbonal area and rounded periphery.
The dorsal and ventral sides have similar convexities (Fig.
21A, C-E, G-J). The adult shell has a diameter of around
1.6mm with 3 whorls and a height of around 0.95mm. The
diameter of the proloculus varies from 0.130 to 0.160µm
(Fig. 21A, E, I, K). The feathering of the intraseptal
interlocular space is marked (Fig. 21C-D, F-H).
Distribution. This species is present in the depositional
sequences Se/Th-1 DS from Andia section (Fig. 9) and in
Lizarraga section (Fig. 10). It aslo occurs in the Th-2 DS
from Korres section (Fig. 13) and in Aixola-Ermua section
(Fig. 14) with the species assemblage indicated in the
aforementioned figures.
Biozone. According to Hottinger (2014) the
biostratigraphic range of this species extends from SBZ 3
to SBZ 6.
Elazigina subsphaerica (sirel, 1972)
Fig. 21K-N

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Paleocene LBF from the Pyrenean Basin

1972 Kathina subsphaerica. Sirel, p. 287; pl. 5, fig.
1-5
1972 Kathina selveri smout. Sirel, p. 290; pl. 5, fig. 6
1983 Kathina aff. subsphaerica sirel. Rahaghi, pl.
36, figs. 17-18
1983 Kathina gr. selveri smout. Rahaghi, pl. 36, figs.
12-15
1988 Kathina selveri smout. Drobne et al., pl. 26, fig.
9
1998 ?Kathina subsphaerica sirel. Accordi et al., p.
200; pl. 16, figs. 1, 3
2004 Smoutina? subsphaerica (sirel). Sirel, p. 12; pl.
8, figs. 1-21
2012 Elazigina subsphaerica sirel, p. 274-275; pl. 2,
figs. 1-17; text fig. 6A-D
2014 Plumokathina subsphaerica (sirel). Hottinger,
p. 117-118; pl. 6.9, figs. 7-16
2015 Elazigina subsphaerica (sirel). Sirel, pl. 30,
figs. 1-9
2018 Elazigina subsphaerica (sirel). Sirel, pl. 8, figs.
1-11
2018 Pseudokathina selveri smout. Sirel, pl. 14, figs.
1-9
Material. Sample Er 5 from Aixola-Ermua section
(Fig. 14).
Description. Large sized specimens with bilamellarperforated test and trochospiral growth. The morphology
is typically subspherical with an angular but unkeeled
periphery. The adult shell has a diameter that varies from
1.3-1.4mm with 2 whorls and height of around 1.20mm.
The diameter of the proloculus is about 0.160-0.170µm in
diameter (Fig. 21K, L).
Distribution. This species has been identified in Th-2
DS from Aixola-Ermua section (Fig. 14) with the species
assemblage indicated in the aforementioned figure.
Biozone.
According
Hottinger
(2014)
the
biostratigraphic range of this species is SBZ 4, from which
it can be considered a marker.
Subfamily Daviesininae hottinger
GENUS Daviesina smout
Type species: Daviesina khatiyahi smout
Daviesina praegarumnensis hottinger, 2014
Fig. 22A-C
2014 Daviesina praegarumnensis n. sp. Hottinger, p.
144; fig. 7.1N-R; pl. 7.18, figs. 1-34
Material. Samples Cm 3 and Cm 6 from Campo section
(Fig. 5) and sample L 3 from Lizarraga section (Fig. 10).

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J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

FIGURE 21. Paleocene Larger Foraminifera from the Pyrenean Basin. A-J: Elazigina lenticula (hottinger, 2014) Specimens A and E from K 2; B, C and
D from sample K 1; F and G from L 26 and H-J from Er 5. K-N: Elazigina subsphaerica (sirel, 1972) Specimens from sample Er 5. Abbreviations: pr:
proloculus, um: umbilical umbo, up: umbilical plate, ilsp: intraseptal interlocular space, spc: spiral canal; fu: funnel, fea: feathering interlocular space

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Description. The test is bilamellar perforate with
trochospiral growth and sub-lenticular morphology. The
dorsal side is decorated with beads and piles, while the
ventral side shows an umbo with funnels. The equatorial
diameter is around 1 mm with 19-20 chambers. The
diameter of the proloculus is around 40µm.
Biozone. The data presented herein validate the
biostratigraphic range SBZ 2-SBZ 3 for this species, as
given by Hottinger (2014).
Daviesina garumnensis
Fig. 22D-E

tambareau,

1972

1972 Daviesina garumnensis. Tambareau, p. 208; pl.
12, figs. 2-5
1980 Daviesina garumnensis tambareau. Caus et al.,
p. 1058; text-fig. 7A-D; pl. 3, figs. 4-6
2014 Daviesina garumnensis tambareau. Hottinger,
p. 144; fig. 7.1N-R; pl. 7.18, figs. 1-34
Material. Sample Se 2 from Serraduy section (Fig. 4),
sample Cm 9 from Campo section (Fig. 5), samples K 1
and K 2 from Korres section (Fig. 13) and sample Er 5
from Aixola-Ermua section (Fig. 14) and Le Quillet and
Cérisols outcrops, located in Petites Pyrénées, France
(Tambareau,1972, v. 1, p. 53).
Description. The test is bilamellar perforate with
lenticular morphology and trochospiral growth. The
spiral growth pattern is alike the one of operculiniform
foraminifera. Beads and piles can be found in dorsal and

ventral sides. The equatorial length is 3.2mm and the
thickness is 1.8mm for 15 chambers. The diameter of the
proloculus of megalospheric forms is about 350µm.
Remarks.
This species is larger than D.
praegarumnensis described by Hottinger (2014), which
he considered a phylogenetic ancestor of D. garumnensis.
Distribution. This species has been identified in
Th-2 DS in Serraduy section (Fig. 4), Campo section (Fig.
5), Korres section (Fig. 13) and Aixola-Ermua section
(Fig. 14) with the species assemblage indicated in the
aforementioned figures.
Biozone. The data presented here validate the
biostratigraphic range of D. garumnensis as SBZ 4, from
which it can be considered a marker, as previously proposed
by Serra-Kiel et al. (1998) and Hottinger (2014).
Superfamily: Planorbulinoidea schwager
Family: Planorbulinida schwager
Subfamily: Planorbulininae schwager
GENUS Planorbulina d’orbigny
Type species: Planorbulina mediterranensis
d’orbigny
T specimens architecturally close to Planorbulina:
wo
P.? antiqua and P.? cretae were identified in the material
studied. The data presented below as well as the observations
from other authors (Hofker, 1966; Hottinger in Peybernès
et al., 2000; Pignatti in Accordi et al., 1998), suggest that
a revision of the architecture of these species is needed in

FIGURE 22. Paleocene Larger Foraminifera from the Pyrenean Basin. A-C: Daviesina praegarumnensis hottinger, 2014. A oblique uncentered equatorial
section; B slightly oblique centred equatorial section; C uncentered axial section, note the funnels in the umbo. Specimens A and B from sample
Cm 6 and C from sample Cm 3. D-E: Daviesina garumnensis tambareau, 1972. D equatorial section; E uncentered axial sections. Specimen D from
Le Quillet outcrop and E from sample Er 5.

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order to clarify their generic attribution. The two species
show significant architectural differences, especially as
regard their ornamentation, which are of generic rank.
Nonetheless, not enough material is available here to carry
out such a revision.
Planorbulina? antiqua
Fig. 23A-H

manguin,

1960

8, L 9 and L 12 from Lizarraga section (Fig. 10), samples
Urb 1, Urb 6, Urb 7, Urb 25, Urb 27 and Urb 28 from
Urbasa Pass section (Fig. 11); sample Le 2 from Leortza
section (Fig. 12); samples Aix 5, Aix 12, Aix 15, Aix
16, Aix 18, Aix 19, Aix 20, and Er 5 from Aixola-Ermua
section (Fig. 14) and samples Ur 6, Ur 13, Ur 14 and Ur 17
from Urko section (Fig. 15).

1960 Planorbulina antiqua. Mangin, p. 275, figs. 44ac
1972 Planorbulina cretae (marsson). Samuel et al.,
pl. 51, fig. 2
1969 Planorbulina cretae (marsson). Bignot and
Larsonneur, pl.3, fig.6

Description. The test is bilamellar-perforate with
flattened morphology. The wall is composed of a thick,
perforate outer layer and a thin, inner layer. Between the
two layers, there is a dark median layer. In axial section,
chambers show an arcuate outline. The proloculus is
spherical with a diameter of around 100-120µm (Fig. 23L).
The apertures are located at the junction of chambers.

Material. Samples Urb 1, Urb 2, Urb 3, Urb 4 from
Urbasa Pass section (Fig. 11) and sample Ur 4 from Urko
section (Fig. 15).

Remarks. P. cretae differs from P. antiqua by the
larger dimensions of the test, by having a thicker wall and
by the presence of beads on the dorsal side.

Description. The test is bilamellar-perforate with
flattened morphology. The wall is composed of a thin,
perforate outer layer and a thin dark inner layer. The chambers
show arcuate outline in axial section. The proloculus is
spherical with a diameter of ca. 40-60µm (Fig. 23E, G). The
apertures are located at the junction of chambers.
Distribution. This species has been identified in the
depositional sequence Ma-Da DS from Urbasa Pass section
(Fig. 11) and in Da-1 DS from Urko section (Fig. 15) with
the species assemblage indicated in the aforementioned
figures.

Distribution. This species is present in the depositional
sequence Ma-Da DS from Urbasa Pass section (Fig. 11). It
also occurs in the Da-1 DS from Lizarraga section (Fig.
10), Urbasa Pass section (Fig. 11), Aixola-Ermua section
(Fig. 14) and in Urko section (Fig. 15); in the Da-2 DS from
Lizarraga section (Fig. 10), Aixola-Ermua section (Fig. 14)
and in Urko section (Fig. 15); in the Se/Th-1 DS from Tena
section (Fig. 6); Andia section (Fig. 9); Lizarraga section
(Fig. 10); Urbasa Pass (Fig. 11); Leortza section (Fig. 12)
and Aixola-Ermua section (Fig. 14); and finally in the Th-2
DS from Aixola-Ermua section (Fig. 14) with the species
assemblage indicated in the aforementioned figures.

Biozone. According to Bignot and Larsonneur (1969)
and to the observations presented here, P.? antiqua has a
biostratigraphic range from Maastrichtian to SBZ 2.

Age. According to Bignot and Larsonneur (1969)
and to the observations presented here P.? cretae has a
biostratigraphic range Maastrichtian-Paleocene.

Planorbulina? cretae (marsson, 1878)
Fig. 23I-M
1969 Planorbulina cretae (marsson). Bignot and
Larsonneur, p. 34, 38; pl. 2, figs. 5-6; pl. 3, figs. 1-6
1972 Planorbulina cretae (marsson). Samuel et al.,
pl. 49, figs. 1-2; pl. 50, figs. 1-2; pl. 51, figs. 1 and 3
1998 Planorbulina cretae (marsson). Sirel, p. 104; pl.
16, figs. 4-12
1998 ?Planorbulina cretae (marsson). Accordi et al.,
pl. 11, fig. 8
2015 Planorbulina cretae (marsson). Sirel, pl. 1, figs.
4-12
2018 Planorbulina cretae (marsson). Sirel, pl. 1, figs.
4-12
Material. Sample Te 2 from Tena section (Fig. 6),
sample An 10 from Andia section (Fig. 9), samples L 2, L

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

ACERVULINOIDEA schultze
Family: Acervulinidae schultze
GENUS Solenomeris douvillé
Type species: Solenomeris ogormani, douvillé
Solenomeris cf. ogormani
Fig. 23N

douvillé,

1924

2003 Solenomeris ogormani douvillé. Bassi, p. 339341; figs. 1 and 2
Material. Samples Urb 1, Urb 3 and Urb 4 from
Urbasa Pass Section (Fig. 11) and sample Le 2 from Leortza
section (Fig. 12).
Remarks. Solenomeris was formerly considered as
belonging to the red algae. Subsequently the studies of
Perrin (1987, 1994) and Bassi (2003) concluded that it

42

J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

FIGURE 23. Paleocene Larger Foraminifera from the Pyrenean Basin. A-H: Planorbulina? antiqua mangin, 1960. A, B, F and G centered axial section;
C and H uncentered axial section; D uncentered longitudinal section and E centered longitudinal section. Specimens A and G from sample Urb 3;
B from sample Urb 4; E from sample Ur 4 and C, D, F and H from sample Urb 1. I-M: Planorbulina? cretae (marsson, 1878) I, L and M subaxial
sections; J and K oblique longitudinal sections. Specimen I from sample Aix 5; J from sample Le 2; K from sample Er 5; L from sample Urb 1 and M
from sample Ur 6. N: Solenomeris cf. ogormani douvillé, 1924 specimen from sample Urb 1. O-Y: Stomatorbina? binkhorsti (reuss, 1862). O, P, Q, R, W
centered axial sections; T centered equatorial section; S, U, V, X and Y uncentered axial sections. Specimen O from sample Ur 6; Q from sample Ur
10; R from sample Urb 3; S from sample Urb 4; P, T, U, V, W and X from sample Urb 1 and Y from sample L 4. Z-C’: Sistanites iranicus rahaghi, 1983
Z equatorial section; A’ and B’ oblique sections and C’ axial section. Specimens Z-B’ from sample L 25 and C’ from sample Eu 2. D’-K’: Valvulineria
patalaensis haque, 1956. D’, E’, F’, G’, H’ equatorial sections; I’ oblique equatorial section and J’-K’ axial sections. Specimen D’ from sample Ur 10; E’,
F’ and I’ from sample Urb 1; G’, H’ and J’ from sample Aix 5 and K’ from sample L 4. L’-R’ Valvulineria bacetai n. sp. L’ Holotype, M’ Paratype; N’ and O’
equatorial sections; P’-Q’ axial sections; M’ and R’ oblique section. Specimens L’ and M’ from sample K 2; N’ from sample Ain 5; O’ from sample Urb
31; P’ from sample L 27 and Q’ and R’ from sample Cm 5. S’-W’: Gavelinellidae indet. S’ centered equatorial section; T’-W’ centered axial sections.
Specimens S’ and U’ from sample Aix 16; T’ from sample Er 5; V’ from sample Beo 5 and W’ from Ur 17. X’-B’’: Coleites cf. reticulosus X’ uncentered
subaxial section Y’ subaxial section, note the carinate margin; Z’ centered oblique axial section, note the spines and A’’ peripheral section, note the
reticulate ornamentations. Specimen X’ from sample Cm 5; Y’, Z’ and B’’ from sample L 15 and A’’ from sample Urb 27.

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J. Serra-Kiel et al.

should be ascribed to the acervulinid foraminifera. We refer
the reader to these studies for further information about the
architecture of this species.
Distribution. The morphology of this species suggests
a habitus attached to the “seagrass” and substrates of reef
environments of the inner shelf. In the material studied this
species was found in the depositional sequence Ma-Da DS
and Se/Th-1 DS from Leortza section (Fig. 12) with the
species assemblage indicated in the aforementioned figures.
Age. According to Perrin (1987, 1994) and Bassi (2003)
this species has a wide biostratigraphic range, exending
from the Paleocene to the Holocene.
Superfamily Discorboidea ehrenberg
Family: Mississippinidae saidova
Subfamily: Stomatorbininae saidova
GENUS Stomatorbina doreen
Type species: Lamarckina torrei cushman and
beermúde

Stomatorbina? binkhorsti (reuss, 1862)
Fig. 23O-Y
1972 Mississippina binkhorsti (reuss). Samuel et al.,
pl. 36, figs. 1-4
1998 Mississippina? binkhorsti (reuss). Sirel, p. 103;
pl. 1, figs. 2, 3, 5, 8-9
1998 Mississippina? sp. Sirel, pl. 1, figs. 1, 5, 7 and 10
1998 Stomatorbina? binkhorsti (reuss). Accordi et
al., pl. 13, fig. 6
2008 Stomatorbina binkhorsti (reuss). Pignatti et al.,
pl. 6, figs. 1-2
2010 Stomatorbina binkhorsti (reuss). Di Carlo et
al., p. 66-67; pl. 1, fig. 11
2015 Stomatorbina binkhorsti (reuss). Sirel, pl. 28,
figs. 6-15
2018 Stomatorbina binkhorsti (reuss). Sirel, pl. 56,
figs. 6-15
Material. samples Sample An 10 from Andia section
(Fig. 9); L 1, L 2, L 4, L 5, L 6 and L 7 from Lizarraga
section (Fig. 10); samples Urb 1, Urb 2, Urb 3, Urb 4, Urb
5, Urb 6, Urb 7, Urb 8, Urb 10, Urb 11, Urb 14, Urb 15,
Urb 17 and Urb 18 from Urbasa Pass section (Fig. 11);
samples Aix 1, Aix 2, Aix 3, Aix 4, Aix 5, Aix 6 and Aix 9
from Aixola-Ermua section (Fig. 14); samples Ur 1, Ur 2,
Ur 3, Ur 4, Ur 5, Ur 6, Ur 7, Ur 9, Ur 10, Ur 13, Ur 14, Ur
16 and Ur 17 from Urko section (Fig. 15).
Description. Low trochospiral test with slightly
biconvex morphology. However, some specimens are
almost flattened. The periphery of the shell is rounded.
The wall is composed of a thick, hyaline outer layer and a

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dark inner layer. For three whorls, the equatorial diameter
is around 1mm. The proloculus is spherical with a diameter
of around 50-80µm (Fig. 23O-Q, S). The dorsal side is
smooth; some specimens show small beads (Fig. 23Q, R).
Septa are inclined backward and the foramina are located in
interiomarginal position (Fig. 23U).
Remarks. Samuel et al. (1972) and Sirel (1998)
determined specimens similar to the specimens studied
here as Mississippina binkhorsti (reuss, 1862). However,
this species has been ascribed to the genus Stomatorbina by
Hottinger in Peybernès et al. (2000), Pignatti in Accordi et
al. (1998) and Sirel (2015). Being aware that the taxonomic
determination of this species remains uncertain, we have
ascribed it tentatively to the genus Stomatorbina.
Distribution. This species is present in the depositional
sequence Ma-Da DS from Urbasa Pass section (Fig. 11), in
Aixola-Ermua section (Fig. 14). It also occurs in the Da-1
DS from Urbasa Pass section (Fig. 11), Aixola-Ermua
section (Fig. 14) and in Urko section (Fig. 15); in the Da-2
DS from Lizarraga section (Fig. 10), Urbasa Pass section
(Fig. 11), Aixola-Ermua section (Fig. 14) and in Urko
section (Fig. 15) with the species assemblage indicated in
the aforementioned figures.
Biozone. The biostratigraphic range of this species is
SBZ 1-SBZ 2.
Sistanites iranicus
Fig. 23Z-C’

rahaghi,

1983

1983 Sistanites iranica. rahaghi, p. 54-55; pl. 34, figs.
1-15
2008 Sistanites iranicus rahaghi. Pignatti et al., pl.
6, fig. 4
2015 Sistanites iranica rahaghi. Sirel, pl. 16, figs.
1-13
2018 Sistanites iranica rahaghi. Sirel, pl. 10, figs.
1-11; pl.11, figs. 1-13.
Material. Sample An 10 from Andia section (Fig. 9);
sample L 25 from Lizarraga section (Fig. 10); samples Aix
19, Aix 21, Er 4 and Er 5 from Aixola-Ermua section (Fig.
14) and sample Eu 2 from Eguaras section (Robador, 2005,
Annex, p. 110-111).
Description. Low-trochospiral test with biconvex
morphology. The periphery of the shell is subacute. The wall
is composed of a thicker, hyaline outer layer and a dark inner
layer. For ten chambers, the equatorial diameter is 0.8mm.
The proloculus is spherical with a diameter of around
100µm (Fig. 23C’). The dorsal and ventral sides ornamented
with beads (Fig. 23C’). Septa are inclined. The foramina are
cribrate, located in interiomarginal position in the septum.

44

J. Serra-Kiel et al.

Remarks. This species differs from S.? binkhorsti in
the more biconvex morphology, larger size of the test and
distribution of stolons in the septum.
Distribution. This species is present in the depositional
Sequence Se/Th-1 DS from Andia section (Fig. 9);
Lizarraga section (Fig. 10), Urbasa Pass section (Fig. 11)
and Aixola-Ermua section (Fig. 14). It also occurs in the
Th-2 DS from Aixola-Ermua section (Fig. 14).
Biozone. The biostratigraphic range of this species is
SBZ 3-SBZ 4.
Family Bagginidae cushman
Subfamily Secovaininae sliter
GENUS Valvulineria cushman
Type species: Valvulineria californica cushman
Valvulineria patalaensis haque, 1956
Fig. 23D’-K’
1956 Valvulineria patalaensis. Haque, p. 162; pl. 12,
fig. 2
Non 2015 Valvulineria aff. patalaensis haque, Sirel,
pl. 10, figs. 1-19
Material. Sample L 4 from Lizarraga section (Fig. 10);
samples Urb 1, Urb 2 and Urb 3 from Urbasa Pass (Fig.
11); samples Aix 5 and Aix 9 from Aixola-Ermua section
(Fig. 14) and sample Ur 10 from Urko section (Fig. 15).
Description. Lamellar perforate specimens with
trochospiral growth and rounded periphery. The chambers
are inflated and increase gradually in size. The dorsal side
is flat to slightly convex; the ventral side has a depressed
umbilicus. The sutures are slightly depressed. The aperture
is simple and located in interiomarginal position. The
diameter of the test for 14-17 chambers varies from 0.460
to 0.650mm and the thickness of adult tests is around
0.370mm. The diameter of the spherical proloculus is
around 45µm.
Distribution. This species occurs in the depositional
Sequence Ma-Da DS from the Urbasa Pass section (Fig.
11); in the Da-1 DS from the Aixola-Ermua section (Fig.
14) and in Urko section (Fig. 15) in the Da-2 DS from
the Lizarraga section (Fig. 10) and Aixola-Ermua section
(Fig. 14) with the species assemblage indicated in the
aforementioned figures.

Paleocene LBF from the Pyrenean Basin

2018 Valvulineria aff. patalaensis
1-19

haque.

Sirel, figs.

Derivation of name. In honour of J.I. Baceta who
studied the Paleocene of the Pyrenean basin.
Holotype. specimen illustrated in Figure 23.38.
Housed in the palaeontological collections of the Museu de
Ciències Naturals de Barcelona, accession number MGB
89845 LP01.01.
Paratype. specimen illustrated in Figure 23.39 is
designated paratype. Housed in the palaeontological
collections of the Museu de Ciències Naturals de Barcelona,
accession number MGB 89845 LP01.02.
Type locality. Located in the Korres section, near
the village of Korres. Sample K 2. Coordinate: N 42º 51’
54.54’’/W 2º 26’ 13.62’’
.
Type level. “Assilina beds” late Thanetian or SBZ 4.
,
Bed of sample K 2 from Korres section.
Material. Samples Cm 3, Cm 4 and Cm 5 from Campo
section (Fig. 5); samples L 25 and L 27 from Lizarraga
section (Fig. 10); sample Urb 31 from Urbasa Pass section
(Fig. 11); sample K 2 from Korres section (Fig. 13) and
sample Ain 5 from Aintzioa section (Appendix, pp. 64-65,
Robador, 2005).
Diagnosis. Wall finely perforated with low trochospiral
growth and rounded periphery. Dimorphism not observed,
probably restricted to early stages of growth. Chambers
inflated and increasing gradually in size as added. Dorsal
side flat; ventral side with depressed umbilicus. Aperture
simple and located in interiomarginal position. Sutures
slightly depressed. Embryo consisting of a spherical
proloculus with a diameter of around 60µm (Fig. 23N’P’). Adult specimens with a diameter generally of around
0.800mm but reaching up to 1mm (Fig. 23M’) for 28
chambers and 3 whorls (Fig. 23O’). Thickness measured
0.441mm.
Remarks. See previous remarks on V. patalaensis
concerning the generic attribution.

Biozone. The biostratigraphic range of this species is
SBZ 1-SBZ 2.

Distribution. This species is present in the depositional
sequence Se/Th-1 DS from Campo section (Fig. 5),
Lizarraga section (Fig. 10) and in Urbasa Pass section (Fig.
11). It also occcurs in the Th-2 DS from Korres section
(Fig. 13). The species assemblage is indicated in the
aforementioned figures.

Valvulineria bacetai n. sp. Serra-Kiel and Vicedo
Fig. 23W-R’

Biozone. The biostratigraphic range of this species is
SBZ 3-SBZ 4.

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J. Serra-Kiel et al.

Superfamily Chilostomelloidea brady
Family Gavelinellidae hofker
Gavelinellidae indet.
Fig. 23S’-W’
1972 Gyroidina subangulosa (plummer, 1926).
Samuel et al., pl. 37, fig. 1-6
Material. Samples Aix 9, Aix 16, Aix 17, Aix 18 and
Er 5 from Aixola-Ermua section (Fig. 14); Ur 2, Ur 3, Ur
6, Ur 8, Ur 10, Ur 11, Ur 13 and Ur 17 from Urko section
(Fig. 15); sample Beo 5 from Beoburu section (Robador,
2005, Annex, p. 112).
Description. The test is bilamellar with coarse
pores and low-trochospiral growth. The dorsal side is
pseudoinvolute, convex and smooth; the ventral side is
slightly convex with an umbo with large beads (Fig. 23V’
,
W’). The lamination is composed of a thicker and perforate
outer layer and a thin, dark inner layer. The proloculus is
spherical with a diameter of 100µm (Fig. 23S’ T’ W’). The
, ,
equatorial diameter is 0.880mm with 20 chambers, and the
thickness 0.490mm.
Distribution. This species is present in the depositional
sequence Da-1 DS from Urko section (Fig. 15). It also
occurs in the Da-2 DS from Aixola-Ermua section (Fig.
14) and in Urko section (Fig. 15), in the Se/Th-1 DS from
Aixola-Ermua section (Fig. 14) and finally in theTh-2 DS
from Aixola-Ermua section (Fig. 14).
Biozone. The biostratigraphic range of this species is
SBZ 1? –SBZ 3.
Family Coleitidae loeblich and tappan
GENUS Coleites plummer
Type species: Pulvinulina reticulosa plummer
Coleites cf. reticulosus (plummer, 1927)
Fig. 23X’-B’’
1973 Coleites sp. aff. C. reticulosus (plummer, 1926).
Ferrer et al. p. 67; fig. 25
1987 Coleites reticulosus (plummer, 1926). Loeblich
and Tappan, pl. 726, fig. 1-3
Material. Sample Cm 5 from Campo section (Fig. 5);
L 15 from Lizarraga section (Fig. 10) and sample Urb 27
from Urbasa Pass section (Fig. 11).
Description. Hyaline test with low-trochospiral
growth (Fig. 23Z’). The chamber sutures are slightly
depressed. In axial section, the periphery is carinate.
The ornamentation consists of long and thin cross ridges

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Paleocene LBF from the Pyrenean Basin

producing a coarsely reticulate pattern in the test surface
(Fig. 23A In axial section, the ridges appear like spines
’’).
(Fig. 23X’ 23Z’).
,
Remarks.
Due to insufficient material the
determination of these specimens cannot be further refined.
Distribution. This species is present in the depositional
sequence Da-2 DS from Lizarraga section (Fig. 10) and
in Urko section (Fig. 15). It also occurs in the Se/Th-1
DS from and Campo section (Fig. 5) and in Urbasa Pass
(Fig. 11) with the species assemblage indicated in the
aforementioned figures.
Biozone. According to the present study, the
biostratigraphic range of this species is SBZ 2-SBZ 3.
Superfamily Nonionoidea schultze
Family Miscellaneidae sigal in piveteau
Subfamily Miscellaneinae sigal in piveteau
GENUS Miscellanea pfender
Type species: Nummulites miscella d’archiac and
haime

Miscellanea yvettae
Fig. 24A-C

leppig,

1988

1988 Miscellanea sp. 2. Drobne et al. p. 160; pl. 26,
figs.1-4
1988a Miscellanea yvettae. Leppig, p. 702; pl. 1, fig. 2;
pl. 2, fig. 2; pl. 3, fig. 2; pl. 5, figs. 1-8
1998 Miscellanea juliettae leppig. Sirel, p. 93; pl. 54,
figs. 1-10
2009 Miscellanea yvettae leppig. Hottinger, p. 5; pl. 7,
figs. 9-16; pl. 8, figs. 1-9; pl. 9, figs. 1-12
Material. Sample Cm 4 from Campo section (Fig. 5);
sample G 3 from Garralda section (Fig. 7) and samples Le 5
and Le 7 from Leortza section (Fig. 12).
Description. The test is bilamellar perforate with
planispiral growth. The lenticular tests are dimorphic
and have a thick wall and rounded periphery. The
ornamentation is dense and it is composed of piles and
pustules. Megalospheric forms show a proloculus with a
diameter of around 200-280µm. The equatorial diameter
in megalospheric forms varies from 1.7 to 1.8mm
and thickness between 1.0-1.15mm for 3 whorls. The
microspheric form starts with a very small proloculus
followed by chambers planispirally coiled, producing an
adult test with an equatorial diameter of 2 mm and axial
thickness of 1.6mm.
Distribution. This species is present in the Se/Th-1
DS in the Campo section (Fig. 5), Garralda section (Fig. 7)

46

FIGURE 24. Paleocene Larger Foraminifera from the Pyrenean Basin. A-C: Miscellanea yvettae leppig, 1988 A axial section, microspheric form; B slightly oblique axial section; C centered axial section;
specimen A from sample Le 5; B from samples Le 7; C from sample Cm 4. D-J: Miscellanea juliettae leppig, 1988 D slightly oblique equatorial section; E equatorial section; F-H axial sections; I-J
subaxial section. Specimens E, F and H from sample Le 5; G from sample Er 1; D, I and J from sample Urb 26. K-P: Miscellanites globularis (rahaghi, 1978) K-M subequatorial sections; N centered
equatorial section and O and P axial sections. Specimen K from sample Ur 6; L and M from sample Ur 3; N from sample Beo 4; O from sample Ur 15 and P from sample L 20. Q-S: Miscellanites
minutus (rahaghi, 1978) equatorial sections, megalospheric forms; specimens Q and R from sample Orio 3 and S from sample G 3. T-X: Miscellanites primitivus (rahaghi, 1983) T subequatorial section;
U-W axial sections; X oblique section. Specimen U from sample Urb 31; T, V, W and X from sample Cm 2. Y-E’: Ornatononion moorkensii hottinger, 2009 Y and Z axial sections. A’-C’ equatorial
sections; D’-E’ tangential sections. Specimens Y and D’ from sample L 22; Z, A’ and E’ from sample Tap 15; B’ and C’ from sample L 21.

J. Serra-Kiel et al.

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J. Serra-Kiel et al.

and in Leortza section (Fig. 12) with the species assemblage
indicated in the aforementioned figures.
Biozone. The stratigraphic data and the species
assemblage confirm the biostratigraphic range given by
Leppig (1988a), Serra-Kiel et al. (1998) and Hottinger
(2009), who considered this species as a marker of the SBZ
3.
Miscellanea juliettae leppig, 1988
Fig. 24D-J
1988a Miscellanea juliettae pfenderae. Leppig, p.
700; pl. 1, fig. 4; pl. 2, fig. 4; pl. 3, figs. 4/1 and 4/2; pl.
4, figs. 1-8, non figs. 9-10
2001 Miscellanea juiettae villattae leppig. Ogorelec
et al.; pl.10, figs. 1-5	
2009 Miscellanea juliettae leppig. Sirel, pl. 4, figs. 7-8
2009 Miscellanea juliettae leppig. Hottinger, p. 6; pl.
1, figs. 21-27; pl. 10, figs. 1-20; pl. 11, figs. 1-11
Material. Samples Urb 26, Urb 29 and Urb 30 from
Urbasa Pass (Fig. 11); samples Le 1, Le 2, Le 4 and Le
5 from Leortza section (Fig. 12) and sample Er 1 from
Aixola-Ermua section (Fig. 14).
Description. Bilamellar-perforate, planispiral test.
The shells are lenticular with thin wall; the periphery is
acute without keel. The ornamentation is composed of
beads covering the surface at the polar zone and abundant
piles covering the lateral surface of the chambers. The
megalospheric forms have an equatorial diameter that varies
from 1.0 to 1.3mm with 3 whorls, and axial thickness of
about 0.7-0.8mm. The megalospheric proloculus is about
160-230µm in diameter.
Distribution. This species is present in Se/Th-1 DS in
the Urbasa Pass section (Fig. 11), the Leortza section (Fig.
12), the Aixola-Ermua section (Fig. 14) and in a sample
taken from a pebble of the breccia located in TH-2 DS.
The species assemblage is indicated in the aforementioned
figures.
Biozone. Stratigraphic data and species assemblage
confirm the biostratigraphic range given by Leppig
(1988a), that leads to consider this species as a marker
of SBZ 3, as proposed by Serra-Kiel et al. (1998) and
Hottinger (2009).
Subfamily Miscellanitinae hottinger
GENUS Miscellanites hottinger
Type species: Miscellanea iranica (rahaghi)
Miscellanites globularis (rahaghi, 1978)
Fig. 24K-P

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1978 Miscellanea globularis. Rahaghi, p. 61; pl. 12,
figs. 10-20
1983 Miscellanea globularis rahaghi. Rahaghi, p. 61;
pl. 42, figs. 1-5 and 7
1998 Miscellanea globularis? rahaghi. Sirel, p. 97; pl.
58; figs. 1-8, 10, 12-14
2000 “Miscellanites” globularis (rahaghi). Peybernès
et al., p. 47; fig. 4a
2009 Miscellanites globularis (rahaghi). Hottinger, p.
10; pl. 22, figs. 1-24; 7-13
2015 Unidentified miscellanid genus. Sirel, pl. 8, figs.
1-7
2018 Burdurina selandinica. Sirel, pl. 19, figs. 1-7
Material. Samples G 1 and G 2 from Garralda section
(Fig. 7); sample An 9 from Andia section (Fig. 9); samples
L 11 and L 20 from Lizarraga section (Fig. 10); samples
Urb 8, Urb 10, Urb 11 and Urb 13 from Urbasa Pass section
(Fig. 11); samples Aix 4, Aix 6, Aix 9, Aix 16 and Aix 17
from Aixola-Ermua section (Fig. 14); samples Ur 3, Ur 5,
Ur 6, Ur 11, Ur 14, Ur 15 and Ur 17 from Urko section
(Fig. 15) and sample Beo 4 from Beoburu section (Robador,
2005, Annex, p. 112).
Description. The test is bilamellar perforate with
planispiral growth. The morphology is spherical to ovoidal.
The surface is covered with pustules. The equatorial
diameter in megalospheric forms varies from 0.6 to 0.8mm
with 3 whorls; the axial thickness is about 0.5mm. There
are 3-4 piles at the polar zones. The protoconch is separated
from the deuteroconch by a thin wall. The proloculus
diameter varies from 70 to 100µm. Chambers have an
isometric outline in equatorial section.
Distribution. This species is present in the depositional
sequence Da-1 DS from Urbasa Pass section (Fig. 11), in
the Aixola-Ermua section (Fig. 14) and in Urko section
(Fig. 15). It also occurs in the Da-2 DS from Garralda
section (Fig. 7), Lizarraga section (Fig. 10), Urbasa Pass
section (Fig. 11), Aixola-Ermua section (Fig. 14) and in
Urko section (Fig. 15). Finally, it has been found in the Se/
Th-1 DS from Andia section (Fig. 9) and in Aixola-Ermua
section (Fig. 14). The species assemblage is indicated in
the aforementioned figures.
Biozone. The stratigraphic data and the associated
foraminifera confirm the biostratigraphic range from SBZ
2 to SBZ 3 attributed by Hottinger (2009) to this species.
Miscellanites minutus (rahaghi, 1978)
Fig. 24Q-S
1983 Miscellanea minuta. rahaghi, p. 62; pl. 43, figs.
1-13
2008 Miscellanea aff. juliettae leppig. Pignatti et al.,

48

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Paleocene LBF from the Pyrenean Basin

p. 134, pl. 6, fig.8-9
2008 Miscellanea cf. juliettae leppig. Pignatti et al., p.
134, pl. 4, fig. 4-5
2009 Miscellanites minutus (rahaghi). Hottinger, p.
8-9; pl. 18, figs. 1-21; pl. 19, figs. 1-11

Distribution. This species is present in Se/Th-1 DS
in the Campo section (Fig. 5), Andia section (Fig. 9),
Lizarraga section (Fig. 10) and in Urbasa Pass section
(Fig. 11) with the species assemblage indicated in the
aforementioned figures.

Material. sample G 3 from Garralda section (Fig. 7);
sample Orio 3 from a pebble taken from Orio section (p.
248, fig. 75, Baceta, 1996) and sample Bur 3 from Burgi
section (Robador, 2005, Annex, pp. 78-79).

Biozone. Stratigraphic data and associated foraminifera
confirm the biostratigraphic range of SBZ 3 attributed to
this species by Hottinger (2009).

Description. The test is bilamellar-perforate with
planispiral growth. The morphology is lenticular. In
equatorial section, chambers have isometric outline. The
proloculus is small (around 40µm in diameter).
Distribution. This species is present in the Garralda
section (Fig. 7) in Se/Th-1 DS with the species assemblage
indicated in the aforementioned figure.
Biozone.
According to Hottinger (2009) the
biostratigraphic range of this species is SBZ 3.
Miscellanites primitivus (rahaghi, 1983)
Fig. 24T
-X
1983 Miscellanea primitiva. Rahaghi, p. 61; pl. 42,
figs. 8-16
1998 Miscellanea sp. 2. Pignatti et al., pl. 4, figs. 8-16
2008 Miscellanea aff. iranica rahaghi. Pignatti et al.,
pl. 6, figs. 5-6
2009 Miscellanites primitivus (rahaghi). Hottinger, p.
8; pl. 16, figs. 1-24; pl. 17, figs. 1-16
2009 Akbarina primitiva (rahaghi). Sirel, p. 419-420;
pl. 3, figs. 1-8
2015 Akbarina primitiva (rahaghi). Sirel, pl. 8, figs.
8-23
2018 Akbarina yarisliensis. Sirel, pl. 19, figs. 8-23
Material. Samples Cm 2, Cm 3 and Cm 4 from
Campo section (Fig. 5); sample An 10 from Andia section
(Fig. 9); samples L 25, L 26 and L 27 from Lizarraga
section (Fig. 10) and sample Urb 31 from Urbasa Pass
section (Fig. 11).
Description. The test is bilamellar perforated with
planispiral growth. The shell is small with lenticular
morphology and rounded periphery. In megalospheric
forms, the equatorial diameter varies from 0.7 to
0.95mm, and axial thickness from 0.45 to 0.50mm. The
ornamentation is composed of thick piles at the polar
zone and fine piles on the lateral surface. The diameter
of the megalospheric forms varies from 60 to 80µm. The
septa are curved and inclined backward at the top of the
chambers.

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

GENUS: Ornatononion hottinger
Type species: Nonion? ornatum van bellen in
moorkens

Ornatononion moorkensii hottinger, 2009
Fig. 24Y
-E’
2009 Ornatononion moorkensii. Hottinger, p. 9-10; pl.
20, figs. 1-13; pl. 21, figs. 1-16
Material. Sample G 1 from Garralda section (Fig. 7);
samples L 12, L 21 and L 22 from Lizarraga section (Fig.
10); samples Urb 8 and Urb 11 from Urbasa Pass section
(Fig. 11); samples Ur 6, Ur 8, Ur 14, Ur 15, Ur 17 and Ur
18 from Urko section (Fig. 15) and sample Tap 15 from
Tapla section (Robador, 2005, Annex, p. 48-49).
Description. The test is bilamellar perforated
with planispiral growth. The morphology of the shell
is lenticular with acute poles. The ornamentation is
composed of pustules and heavy piles in the polar zones.
The megalospheric forms have an equatorial diameter that
varies from 1.2 to 1.3mm, and axial thickness of around
0.6mm. The diameter of the proloculus of megalospheric
forms is around 100µm.
Distribution. This species is present in the depositional
sequences Da-1 DS from Urbasa Pass section (Fig. 11) and
in Urko section (Fig. 15). It also occurs in the Da-2 DS
from Garralda section (Fig. 7), Lizarraga section (Fig. 10)
and Urko section (Fig. 15).
Biozone. The stratigraphic data and the foraminifera
associated with this species validate its biostratigraphic
attribution to SBZ 2 as given by Hottinger (2009).
Superfamily Nummulitoidea de blainville
Family Nummulitidae de blainville
GENUS: Nummulites lamarck
Type species: Camerina laevigata bruguière
“Operculina” heberti
Nummulites heberti
Tosquella, 1995]
Fig. 25J-O

munier-chalmas,

1884 [=
(munier-chalmas) sensu

49

J. Serra-Kiel et al.

1955 Operculina heberti munier-chalmas. Mangin, p.
241; pl. 3, figs. 1-8
1962 Operculina heberti munier-chalmas. Villatte, p.
291; pl. 21, figs. 1-2; pl. 22, figs. 3-4
1964 Operculina heberti munier-chalmas. Hottinger,
pl. 1, figs. 7a-e
1972 Operculina
heberti
munier-chalmas.
Tambareau, p. 221; pl. 13, figs. 1-2; pl. 14, figs. 3-4
1977 Operculina heberti munier-chalmas. Hottinger,
p. 71; text-figs. 27M-O, 28
1995 Nummulites
heberti (munier-chalmas).
Tosquella, p. 293; pl. 2, figs. 1-9
1998b Nummulites
heberti (munier-chalmas).
Tosquella and Serra-Kiel, p. 95-96; pl. 17, figs. 1-3
2008 “Operculina”
heberti (munier-chalmas).
Pignatti et al., pl. 2, fig. 2
Material. Sample Te 2 from Tena section (Fig. 6);
samples Urrob 3 and Urrob 4 from Urrobi section (Fig. 8);
samples An 1, An 2 and An 11 from Andia section (Fig. 9);
samples L 24, L 25 and L 26 from Lizarraga section (Fig.
10); samples Urb 22, Urb 23 and Urb 25 from Urbasa Pass
section (Fig. 11); sample Le 3 from Leortza section (Fig. 12);
samples Aix 15, Aix 16, Aix 17, Aix 18, Aix 21, Er 1 and Er
2 from Aixola-Ermua section (Fig. 14); sample from Bouzin
outcrop located in the Petites Pyrénées (north Pyrenean
realm) see Villatte (1962), Tambareau et al. (1992b, p. 14)
and Schaub (1981, p 193, fig. 45) and sample Narp outcrop
from Béarn (France), see Tambareau et al. (1994a, p. 36).
Description. The morphology of the microspheric
and megalospheric form is flattened. The microspheric
forms have a maximum equatorial diameter of ca. 1mm,
while the megalospheric are around 0.6mm across. The
spire is operculiniform, slightly irregular, opening rapidly.
The ornamentation is composed of piles over the septa
and between them. Diameter proloculus in megalospheric
forms varies from 200 to 250µm. The septa are straight with
dense distribution and recurved in the top of the chambers.
Remarks. This species was originally accommodated
within genus Operculina. However, according to Tosquella
(1995), a trabecular system can be observed in exceptionally
well-preserved specimens from Narp, France (Tambareau
et al., 1994a, p. 36), concluding that the species should
be ranged in genus Nummulites. This feature is again
illustrated here (Fig. 25.15).
However, by transferring the species to Nummulites, it
becomes a junior secondary homonym of Nummulites heberti
d’ rchiac and Haime, 1853. As long as this controversy is not
A
solved, the morphotype should be left in open nomenclature.
Distribution. This species is present in Se/Th-1 DS
in the Tena section (Fig. 6), Urrobi section (Fig. 7), Andia

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

Paleocene LBF from the Pyrenean Basin

section (Fig. 9), Lizarraga section (Fig. 10); Urbasa Pass
section (Fig. 11); Leortza section (Fig. 12) and in AixolaErmua section (Fig. 14), in this section the pebbles taken
from a breccia in Th-2 DS this species is associated with the
species assemblage indicated in the aforementioned figures.
Biozone. The stratigraphic data and the associated
foraminifera confirm the biostratigraphic range SBZ 3
attributed by Tosquella (1995), Tosquella and Serra-Kiel
(1998b) and Serra-Kiel et al. (1998) to this species.
Nummulites catari tosquella and serra-kiel, 1998
Fig. 25P-U
1998a Nummulites catari. Tosquella and Serra-Kiel, p.
165-171; fig. 1; pl. 1, figs. 1-2; pl. 2, figs. 1-4
1998b Nummulites catari tosquella and serra-kiel.
Tosquella and Serra-Kiel, p. 96-97; pl. 17, figs. 4-5
Material. Samples from Cérisols and Le Quillet
outcrops located in Petites Pyrénées (France) as reported
by Tambareau (1972; v.1, p. 53), Tambareau et al. (1992b,
p. 14) and Schaub (1981, p 193 and fig. 45).
Description. The morphology of the microspheric
forms is flattened with rounded periphery and slightly
inflated at the polar regions. The test shows a diameter of
6.55mm for 5 whorls. The ornamentation consists of curved
filaments and small piles over them in the polar zone. The
spire is operculiniform tending to irregular assilinoid. The
megalospheric forms show a flattened morphology, with
a diameter of 3.3-3.5mm and thickness of around 1 mm
for 3 whorls. The spire is regular and opens rapidly. Both
generations show a thin marginal cord, and dense septa
distribution. Septa straight and strongly recurved in the upper
part of the chamber. The proloculus is small in megalospheric
forms, with a diameter of around 120-150µm.
Distribution. In the Le Quillet outcrop this species is
associated with A. yvettae and A. azilensis, while in the
Cérisols outcrop it is associated with A. azilensis.
Biozone. According to Tosquella and Serra-Kiel
(1998b) and Serra-Kiel et al. (1998) its biostratigraphic
range is SBZ 4.
GENUS Assilina D’orbigny
Type species: Assilina depressa, D’orbigny
Assilina yvettae schaub, 1981
Fig. 25A-C
1972 Assilina? sp., Tambareau, p. 217; pl. 13, figs. 9-16;
pl. 14, figs. 16-19; pl. 18, figs. 1-2, 4
1981 Assilina yvettae. Schaub, p. 193; pl. 70, figs.

50

J. Serra-Kiel et al.

1-13; tab. 16, fig. 1.3
1995 Assilina yvettae schaub. Tosquella, p. 310; pl.
42, figs. 1-4
1998b Assilina yvettae schaub. Tosquella and SerraKiel, p. 101-102; pl. 18, figs. 1-3
2015 Assilina yvettae schaub. Sirel, pl. 23, figs. 1-12
2018 Assilina yvettae schaub. Sirel, pl. 25, figs. 1-17
Material. Sample from Le Quillet outcrop located
in the Petites Pyrénées, see Tambareau (1972, v. 1, p. 53)
and Schaub (1991, p 193 and fig. 45); sample Cm 9 from
Campo section (Fig. 5); sample Te 6 from Tena section (Fig.
6); samples Urrob 6 and Urrob 7 from Urrobi section (Fig.
8); samples K 1 and K 2 from Korres section (Fig. 13) and
sample Er 5 from Aixola-Ermua section (Fig. 14).
Description. Hyaline test with planispiral, evolute
coiling. Microspheric forms not found. The morphology of
the test is lenticular-flattened with a rounded periphery. The
equatorial diameter is 2.7mm and the thickness is 1.1mm
for 4-5 whorls. The spire is regular and closed, assilinoid
type, and the marginal cord is thin. The ornamentation is
composed of piles over the filaments and at the polar zone.
The septa are regularly distributed and straight, slightly
curved backward at the top of the chambers. In equatorial
section, the outline of the chambers is subrectangular and
is higher than wide. The diameter of the proloculus is 100150µm.
Distribution. This species is identified in Th-2 DS in
the following sections: Campo section (Fig. 5); Tena section
(Fig. 6); Urrobi section (Fig. 8); Korres section (Fig. 13)
and in Aixola-Ermua section (Fig. 14) with the species
assemblage indicated in the aforementioned figures. In the
Le Quillet outcrop this species is associated with N. catari
and A. azilensis.
Biozone. The stratigraphic data and the associated
foraminiofera show that this species belongs to SBZ 4,
as pointed out by Schaub (1981) and Serra-Kiel et al.
(1998).
Assilina azilensis (tambareau, 1966)
Fig. 25D-I
1964 Operculina aff. douvillei doncieux. Hottinger,
pl. 1, figs. 5a-e, figs. 10-13
1966 Operculina (Nummulitoides) azilensis.
Tambareau, p. 301, pl. 1, figs. 1-9
1969 Ranikothalia azilensis (tambareau). Butterlin
and Monod, p. 600; pl. 1, figs. 5, 6 and 9
1977 Operculina azilensis tambareau. Hottinger, p.
63, text-figs. 22N-S; pl. 23, figs. 1-10
1995 Assilina azilensis (tambareau). Tosquella, p.
351; pl. 42, figs. 1-5

Geologica Acta, 18.8, 1-69, I-III (2020)
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Paleocene LBF from the Pyrenean Basin

1998b Assilina azilensis (tambareau). Tosquella and
Serra-Kiel, p. 118-119; pl. 23, figs. 7-9
Material. Samples from Cérisols and Le Quillet
outcrops located in Petites Pyrénées (France), see Tambareau
(1972) and Tambareau et al. (1992b, p.14); sample Cm 9
and Cm 10 from Campo section (Fig. 5); sample Te 6 from
Tena section (Fig. 6); samples Urrob 6 and Urrob 7 from
Urrobi section (Fig. 8); samples Urb 32 and Urb 33 from
Urbasa Pass section (Fig. 11); samples Le 9 and Le 10 from
Leortza section (Fig. 12); sample Er 5 from Aixola-Ermua
section (Fig. 14).
Description. The tests of the micro- and megalospheric
forms show a flattened morphology, slightly inflate at the
polar region and have a rounded periphery. The growth
is evolute. The microspheric forms show an equatorial
diameter of around 5.1 mm for 5 whorls and a thickness
around 0.75mm in axial section. The megalospheric forms
have an equatorial diameter of around 3.6mm for 3 whorls
and a thickness around 0.7-1.1mm in axial section. In both
generations, the spire is regular and opens rapidly. Septa
are straight or slightly inclined and curved backward at the
top of the chambers. The ornamentation consists of piles
concentred at the polar zone. In megalospheric forms, the
embryonic apparatus is isolepidine type. The diameter of
the protoconch is 190-240µm and the deuteroconch has a
length of 170-200µm in axial section and a height of 110160µm.
Distribution. This species is identified in Th-2 DS
in the following sections: Campo section (Fig. 5); Tena
section (Fig. 6); Urrobi section (Fig. 8); Urbasa Pass section
(Fig. 11); Leortza section (Fig. 12) and in Aixola-Ermua
section (Fig. 14) with the species assemblage indicated in
the aforementioned figures. In the Cérisols outcrop it is
associated with N. catari and in the Le Quillet outcrop
with A. yvettae and N. catari.
Biozone. According to the stratigraphic data available
and the associated foraminifera, the species belongs to SBZ
4, as pointed out by Hottinger (1977) and Serra-Kiel et al.
(1998).
GENUS Ranikothalia caudri
Type species: Nummulites nuttalli davies
Ranikothalia soldadensis (vaughan
1941)
Figs. 25V-B’ 26A-M
,

and cole,

1941 Miscellanea soldadensis. Vaughan and Cole, p.
36; pl. 4, figs. 8-9
1944 Ranikothalia soldadensis (vaughan and cole).
Caudri, p. 23; pl. 4, fig. 19; pl. 5, figs. 24, 26?

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J. Serra-Kiel et al.

Paleocene LBF from the Pyrenean Basin

FIGURE 25. Paleocene Larger Foraminifera from the Pyrenean Basin. A-C: Assilina yvettae schaub, 1981, A centered equatorial section; B external
view and C axial section; all specimens megalospheric forms. Specimens A and B from sample Le Quillet outcrop and C from sample K 2. D-I: Assilina
azilensis (tambareau, 1966), D, F and G centered equatorial sections, megalospheric forms; E external view, megalospheric form; H centered equatorial
section, microspheric form and I axial section. All specimens from sample Cérisols outcrop. J-O: “Operculina” heberti (munier-chalmas, 1884), J
external view, megalospheric form; K-N centered equatorial sections, megalospheric forms; O oblique equatorial section, megalospheric form, note
the trabecular system. Specimens J-N from sample Bouzin outcrop and 15 from Narp outcrop. P-U: Nummulites catari tosquella and serra-kiel, 1998,
P centered equatorial section, microspheric form; Q-S centered equatorial sections, megalospheric forms; T external view, megalospheric form and
U axial section, megalospheric form. All specimens from sample from Cérisols outcrop. V-B’: Ranikothalia soldadensis (vaughan and cole, 1941), V
equatorial section, microspheric form; W external view, microspheric section; X and Z centered equatorial sections, megalospheric forms; Y external
view, megalospheric form; A’ centered axial section, megalospheric form; B’ axial section, microspheric form. Specimens V-Z from sample Ruisseau
de la Mède outcrop; A’ from sample Aix 14 and B’ from Er 1. Abbreviations: tr: trabecular system

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J. Serra-Kiel et al.

1945 Miscellanea soldadensis (vaughan and cole).
Vaughan, p. 30; pl. 5, figs. 2-5
1960 Operculina catenula (cushman and jarvis).
Butterlin and Bonet, p. 11-14; pl. 2, figs. 1-5
1975 Ranikothalia soldadensis (vaughan and cole).
Caudri, p. 539; pl. 1, figs. 1, 4; pl. 2, figs. 1, 6, 8; pl. 6,
figs. 1, 3; pl. 7, figs. 1-5, pl. 8, figs. 1-3
1996 Ranikothalia soldadensis (vaughan and cole).
Caudri, p. 1185; pl. 5.4; pl. 9.1
1997 Ranikothalia soldadensis (vaughan and cole).
Berlanga, p. 145-151; pl. 30, figs. 1-4; pl. 31, figs. 1-3;
pl. 32, figs. 1-6; pl. 33, fig. 1
2014 Ranikothalia soldadensis (vaughan and cole).
Vicedo et al., p. 58-62; figs.14-16
Material. Sample from Ruisseau de la Mède outcrop
located in Plantaurel region, north Pyrenean realm, see
Villatte (1962) and Tambareau (1972); samples Urb 26 and
Urb 28 from Urbasa Pass section (Fig. 11) and samples Aix
13, Aix 14 and Er 1 from Aixola-Ermua section (Fig. 14).
Description. The microspheric forms show a flat
morphology. The equatorial diameter is around 7.2mm
and the axial thickness is around 1.3mm for 4 whorls.
The megalospheric forms show a lenticular-flattened
morphology with rounded periphery. In both generations,
the growth is involute, becoming evolute in the last whorls.
The ornamentation is composed of septal filaments and
large piles at the polar zone. The septa are slightly curved.
The marginal cord is thick. The proloculus in megalospheric
forms has a diameter varying from 150 to 180µm.
Remarks. Comparison of specimens of R. sindensis from
the Paleocene of the Salt Range (Pakistan), those described
here (Fig. 26P-Q) and those from Libya illustrated by Hottinger
(1977), show that Ranikothalia sindensis differs from
Ranikothalia soldadensis in the larger shell dimensions, the
looser spiral, the larger diameter of the proloculus and the thinner
marginal cord. Inconsistences related to genus Ranikothalia
are discussed in more detail in Vicedo et al. (2014).
Distribution. This species is identified in Se/Th-1
DS in the following section: Urbasa Pass section (Fig.
11) and in Aixola-Ermua section (Fig. 14) where, in the
the pebbles taken from a breccia in Th-2 DS, this species
is associated with the species assemblage indicated in the
aforementioned figures.
Biozone. According the stratigraphic data and the
associated foraminifera this species belongs to SBZ 3, as
pointed out by Tosquella (1995) and Serra-Kiel et al. (1998).
Family Discocyclinidae galloway
GENUS: Discocyclina gümbel
Type species: Orbitolites pratti michelin

Geologica Acta, 18.8, 1-69, I-III (2020)
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Paleocene LBF from the Pyrenean Basin

Discocyclina seunesi
Fig. 27A-B

douville,

1922

1972 Discocyclina seunesi douville. Neumann, pl. 4;
fig. 8
1972 Discocyclina seunesi douville. Samuel et al., p.
159; figs. 1-7.
1987 Orbitocypeus seunesi (douville). Less, p. 194196; text-fig. 30a
2009 Discocyclina seunesi douvillé. Sirel, pl. 4, fig. 5
Material. Sample Te 2 from Tena section (Fig. 6);
sample Urrob 4 from Urrobi section (Fig. 8); samples An
1, An 2, An 5, An 9 and An 11 from Andia section (Fig.
9); samples L 24, L 25 and L 26 from Lizarraga section
(Fig. 10); samples Urb 23, Urb 24, Urb 25 and Urb 27
from Urbasa Pass section (Fig. 11) and sample Le 3 from
Leortza section (Fig. 12).
Description. Small, lamellar perforate test with
lenticular morphology. The equatorial diameter is about
1mm for 30-35 annuli and the thickness about 0.3mm. The
embryonic apparatus is composed of a small protoconch
and a deuteroconch. The diameter of the protoconch is
around 0.1mm. The equatorial chambers increase in size
towards the periphery.
Distribution. This species is present in Se/Th-1 DS in
the following sections: Tena section (Fig. 6); Urrobi section
(Fig. 8); Andia section (Fig. 9); Lizarraga section (Fig.
10); Urbasa Pass section (Fig. 11)and in Leortza section
(Fig. 12) with the species assemblage indicated in the
aforementioned figures.
Biozone. According to Less (1987) this species is
probably Thanetian in age. Later, Less in Serra-Kiel et al.
(1998) stablished its biostratigraphic range as SBZ 3. Our
data confirm this attribution.
Family Pararotaliidae

reiss

Description. Hyaline, trochospiral shells with single or
multiple foramina and a “toothplate” as umbilical closure
of the spiral chamber, with or without a dense enveloping
canal system.
Remarks. This definition of the family is in agreement
with Hottinger’s (2014) point of view that some rotaliform
morphotypes, particularly the representatives of subfamilies
Pararotaliinae reiss, 1963 and Laffiteininae hottinger,
2014 (=Cuvillierininae loeblich and tappan, 1964) should
be excluded from from the Family Rotaliidae. However
Hottinger (2014) in his “Identification Key to Some Genera
Excluded from the Rotaliacea” also proposed to exclude
them from the superfamily Rotalioidea ehrenberg, a

53

J. Serra-Kiel et al.

view that we have not followed here, taking into account
the definition of this taxon given by Loeblich and Tappan
(1987). In consequence the former subfamily Pararotaliinae
is elevated here to the rank of Family Pararotaliidae.
Subfamily Pararotaliinae reiss
GENUS Paralockhartia hottinger
Type species: Paralockhartia eos, hottinger

Paleocene LBF from the Pyrenean Basin

Paralockhartia eos hottinger, 2014
Fig. 27C-D
2014 Paralockhartia eos. Hottinger, p. 157; pl. 8.3,
figs. 1-14; pl. 8.4, figs. 1-15
Material. Sample L 3 from Lizarraga section (Fig. 10)
and sample Ur 18 from Urko section (Fig. 15).

FIGURE 26. Paleocene Larger Foraminifera from the Pyrenean Basin. A-M: Ranikothalia soldadensis (vaughan and cole, 1941) A-E Specimens from
Campeche drawings after Berlanga (1997); F-G Specimens from Dor M’Said drawings after Hottinger (1977). H-I Specimens from Ruisseau de la
Mède drawings after Hottinger (1997). J-M Specimens from Ruisseau de la Mède drawings after our material. N-Q: Ranikothalia sindensis (davies,
1927), specimens from Salt Range (Pakistan) drawings after our material.

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J. Serra-Kiel et al.

Description. The test is bilamellar perforate with trochospiral
growth and lenticular morphology The dorsal and ventral sides
.
have similar convexities. The equatorial diameter is around 1.2mm
for three whorls. The ventral side shows piles and discontinuous
funnels. The proloculus has a diameter of around 60µm.
Distribution. This species is found in Da-2 DS in the
following sections: Lizarraga section (Fig. 10) and in Urko

Paleocene LBF from the Pyrenean Basin

section (Fig. 15) with the species assemblage indicated in
the aforementioned figures.
Biozone. The stratigraphic data and the associated
taxa presented here validate the attribution of this species
to SBZ 2 as proposed by Hottinger (2014).
Subfamily Cuvillierininae loeblich and tappan

FIGURE 27. Paleocene Larger Foraminifera from the Pyrenean Basin. A-B: Discocyclina seunesi douville, 1922. A centred equatorial section; B axial
section. Specimens A and B from sample An 1. C-D: Paralockhartia eos hottinger, 2014 axial sections. Specimen C from sample L 3 and D from
sample Ur 18. E-H: Cuvillierina sireli inan, 1988; E oblique equatorial section; F centered axial section; G and H subaxial sections. Specimen E and
F from sample Urb 26; G from sample Tap 10 and H from sample G 3. I-M: Thalmannita madrugaensis (cushman and bermúdez, 1947). I-K subaxial
sections; L subequatorial section and M centered equatorial section. Specimen I from sample Aix 12; J from sample Le 5; K from sample Urb 25; L
from sample Urrob 3; M from sample Bur 3. N-P: Scarificatina reinholdi (pozaryska and szczechura, 1970). N axial section; O and P subaxial sections.
Specimen N from sample L 12; O from sample Ur 10 and P from sample L 19. Q-W: Cincoriola ovoidea (haque, 1958). Q-V axial sections; W oblique
section. Specimens Q, R, U and W from sample Cm 5; S and T from sample Cm 3 and V from sample Beo 2.

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J. Serra-Kiel et al.

The subfamily Laffitteininae introduced by Hottinger
(2014) is a junior synonym of Cuvillierininae loeblich
and tappan, 1964. As stated by Hottinger (2014, p. 153)
“This group merits to be ranked as a particular subfamily
Cuvillierininae loeblich and tappan, 1964”
.
Genus Cuvillierina debourle
Type species: Laffitteina vallensis, ruiz de gaona
Cuvillierina sireli inan, 1988
Fig. 27E-H
1988 Cuvillierina sireli. Inan, p. 121; pl. 1, figs.1-9; pl.
2, figs. 1-8
1998 Pseudocuvillierina sireli (inan). Sirel, p. 86; pl.
44, figs.1-20
2004 Pseudocuvillierina sireli (inan). Sirel, p. 9; pl.
7, figs. 1-20
2000 “Cuvillierina” sireli inan. Peybernès et al., p. 46;
figs. 6.11-6.3
2009 Pseudocuvillierina sireli (inan). Sirel, pl. 3, figs.
9-13
2014 Cuvillierina sireli inan. Hottinger, p. 164, 166;
pl. 8.7, figs. 1-11; pl. 8.8, figs. 1-12
2008 Pseudocuvillierina sireli (inan). Pignatti et al.,
pl. 6, fig. 11
2010 Pseudocuvillierina sireli (inan). Di Carlo et al.,
p. 70; pl. 1, fig. 12
2015 Pseudocuvillierina sireli (inan). Sirel, pl. 7, figs.
1-21
2018 Pseudocuvillierina sireli (inan). Sirel, pl. 21,
figs. 1-21
Material. sample G 3 from Garralda section (Fig.
7); sample Urb 26 from Urbasa Pass section (Fig. 11);
samples Le 4 and Le 5 from Leortza section (Fig. 12);
samples Er 1, Er 2 and Er 3 from Aixola-Ermua section
(Fig. 14); sample Ur 14 and Ur 17 from Urko section
(Fig. 15) and sample Tap 10 from Tapla section (Robador,
2005, Annex, p. 48-49).
Description. The test is lenticular with a bilamellar
perforate wall. The chamber arrangement is very lowtrochospiral, almost planispiral. The enveloping canal
system covers both sides of the test (Fig. 27G). The
periphery is acute. Both umbos are perforated by funnels.
The equatorial diameter varies from 1.15 to 1.2mm and the
thickness from 0.7 to 0.8mm with two whorls. The diameter
of the proloculus is about 90µm.
Distribution. This species is identified in the
following Depositional Sequences: Da-2 DS from Urko
section (Fig. 15). Se/Th-1 DS from Garralda section (Fig.
7); Urbasa Pass section (Fig. 11); Leortza section (Fig. 12)
and in Aixola-Ermua section (Fig. 14) in two pebbles of the

Geologica Acta, 18.8, 1-69, I-III (2020)
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Paleocene LBF from the Pyrenean Basin

breccia located in Th-2 DS with the species assemblage
indicated in the aforementioned figures.
Biozone. According to Sirel (1998), the age of this
species is Thanetian. Serra-Kiel et al. (1998) considered its
biostratigraphic as ranging from SBZ 2 to the middle part
SBZ 4. The biostratigraphic range of this species from the
middle part SBZ 5 to middle part SBZ 9 illustrated in figure
3.1 of Hottinger (2014) is certainly a mistake introduced
while finishing this book after the author’s decease. In the
study area this species has been found to range from the
middle part of the SBZ 2 to SBZ 3.
Rotaliid taxa of uncertain affinities sensu Hottinger (2014)
Genus Thalmannita bermúdez
Type species: Rotalia madrugaensis, cushman and
bermúdez

Thalmannita
madrugaensis
bermúdez, 1947)
Figs. 27I-27M

(cushman

and

2000 Thalmannita madrugaensis (cushman and
bermúdez), Peybernès et al., p. 46; fig. 4b
2009 Thalmannita madrugaensis (cushman and
bermúdez). Sirel, pl. 1, figs. 1-7
2014 Thalmannita madrugaensis (cushman and
bermúdez). Hottinger, p. 173, 176; pl. 9.1, figs. 1-21
2018 Thalmannita madrugaensis (cushman and
bermúdez). Sirel, pl. 7, figs. 15-22
Material. Sample Te 5 from Tena section (Fig. 5);
sample Urrob 5 from Urrobi section (Fig. 8); sample Urb
25 from Urbasa Pass section (Fig. 11); samples Le 5 and Le
8 from Leortza section (Fig. 12); samples Aix 12, Aix 18,
Aix 21, Er 1 and Er 2 from Aixola-Ermua section (Fig. 14);
sample Bur 3 from Burgi section (Robador, 2005, Annex,
pp. 78-79) and Orio 3 sample from a pebble taken from
Orio section (p. 248, fig. 75, Baceta, 1996).
Description. Bilamellar perforate test with planispiral
growth. The chambers are inflated and the periphery of the
shell is rounded. The ornamentation is formed by abundant
spines. The equatorial diameter is around 0.6mm.
Distribution. This species is identified in Se/Th-1 DS in
the following sections: T section (Fig. 5); Urrobi section (Fig.
ena
8); Urbasa Pass section (Fig. 11); Leortza section (Fig. 12) and
in Aixola-Ermua section (Fig. 14) and in two pebbles collected
in the breccia in DS Th-2 this species is associated with the
species assemblage indicated in the aforementioned figures.
Biozone. According to Serra-Kiel et al. (1998) and
Hottinger (2014), the biostratigraphic range of this species
extends from SBZ 3 to SBZ 4.

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Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

Material. Samples Cm 3 and Cm 5 from Campo
section (Fig. 5) and Sample Beo 2 from Beoburu section
(Robador, 2005, Annex, p. 112).

GENUS Scarificatina moorkens
Type species: Boldia reinholdi, pozaryska and
szczechura

Scarificatina
reinholdi
szczechura, 1970)
Fig. 27N-P

(pozaryska

and

2000 Scarificatina sp., Peybernès et al. p. 46; fig. 6.9
2012 Haymanina danica. Sirel, p. 274; pl. 1, figs. 1-3
2014 Scarificatina
reinholdi (pozaryska and
szczechura). Hottinger, p. 181; pl. 9.3
2015 Haymanina danica. Sirel, pl. 1, figs. 1-3
Material. Samples L 12 and L 19 from Lizarraga
section (Fig. 10); samples Urb 7, Urb 13 from Urbasa Pass
section (Fig. 11); sample Aix 6 from Aixola-Ermua section
(Fig. 14) and samples Ur 10, Ur 11 and Ur 14 from Urko
section (Fig. 15).

Description. Lamellar perforate test with subglobular
morphology and trochospiral growth. The diameter of the
proloculus is around 0.05mm. The dorsal side is flattened,
the ventral side is markedly convex with rounded outline.
For two whorls the equatorial diameter is around 0.6mm0.7mm.
Distribution. This species is present in Campo section
(Fig. 5) in Se/Th-1 DS with the species assemblage
indicated in the aforementioned figures.
Biozone. The data obtained in this study validate the
belonging of this species to the SBZ 3, as pointed out by
Hottinger (2014).
Biostratigraphy based on Larger Foraminifera

Description. The test is bilamellar perforate with
planispiral growth and non-symmetric equatorial
morphology. The equatorial diameter varies from 0.5
to 0.6mm. The wall shows coarse pores. The dorsal
side is f lattened to slightly concave. The ventral side
has parallel crests. The diameter of the proloculus is
around 60µm.
Distribution. This species is present in the following
Depositional Sequences: Da-1 DS from Urbasa Pass
section (Fig. 11); Aixola-Ermua section (Fig. 14 and in
Urko section (Fig. 15). Da-2 DS from Lizarraga section
(Fig. 10); Urbasa Pass section (Fig. 11) and in Urko section
(Fig. 15) with the species assemblage indicated in the
aforementioned figures.
Biozone. According to Hottinger (2014) the
biostratigraphic range of this species extends from the
upper part of SBZ 2 to SBZ 5.
GENUS Cincoriola haque
Type species: Punjabia ovoidea
Cincoriola ovoidea (haque, 1956)
Fig. 27Q-W
1998? Kathina melona (rahaghi). Accordi et al., p.
174; pl. 3, fig. 3.c; p. 196; pl. 14, figs. 1-3
1998 Soriella cf. bitlisica sirel. Accordi et al., p. 190;
pl. 11, fig. 1
2014 Cincoriola ovoidea (haque). Hottinger, p. 183;
pl. 9.4, figs. 14-23; pl. 9.5; 1-14
2018 Soriella bitlisica sirel. Sirel, pl. 7, figs. 7-14
2018 Cincoriella paleocenica. Sirel, pl. 10, figs.
12-14

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

Sixty species of larger foraminifera were found in the
Pyrenean Paleocene with a biostratigraphic distribution
belonging to the Tethyan SBZ 1 to SBZ 4. The new data led
to a revision of the definition of these biozones, as proposed
below.
Remarks on SBZ 1. According Serra Kiel et al.
(1998), this biozone is characterized by the presence
of Laffitteina bibensis and Bangiana hanseni as taxa
defining the beginning of the Danian stage. According to
our database, the use of these two species as markers need
emendation.
Laffitteina bibensis is found in the Campo section
together with the charophytes Peckichara llobregatensis
and Sphaerochara edda attributed to the base of Paleocene
(Fig. 5). Marie (1946) characterized these species from the
Mont Aimé marls (Paris Basin) as being of Montian age.
Later, Bignot (1987), Bignot (1993) and Bignot et al. (1997)
also attributed the Vertus-Mont Aimé Fm. with Laffitteina
bibensis (type locality) to the early Danian. However, the
Paleocene age of the beds containing Laffitteina bibensis
in Campo (sample Cm 1, Fig. 5) was first questioned
by López-Martínez et al. (2006), who pointed out that
these deposits underlie beds with rudists, the ammonite
Pachydiscus gollevillensis and dinosaur remains. Finally,
Guernet and Villier (2017), while studying the ostracods
associated with - in the Mont Aimé Fm., considered that
they were exclusively Cretaceous in age. Altogether, these
data indicate that the attribution of the Laffitteina bibensis
to the Danian should be rejected.
Bangiana hanseni was first described by Drobne et al.
(2007) from the Danian of Slovenia. In the Pyrenees, this

57

J. Serra-Kiel et al.

species has been identified in the sections of Tena (sample
Te 1; Fig. 6), Urrobi (samples Urrob 1 and Urrob 2; Fig. 8)
and Urbasa Pass (samples Urb 2, Urb 12, Urb 16 and Urb
21; Fig. 11). In the Tena and Urrobi sections, this species
is located in the Da-1 DS, while in the Urbasa section (Fig.
11), it is located in the Ma-Da Ds and Da-2 DS. In the
Ma-Da DS Bangiana hanseni occurs associated with P.?
antiqua, and Stomatorbina? binkhorsti, whereas in the
Da-2 DS it is associated with Haymanella paleocenica,
Haymanella elongata and Goesella sp. and overlying the
beds with Miscellanites globularis and Ornatononion
moorkensii indicating an SBZ 2 age.
In the Urbasa Pass section, other species found in the
lower and middle parts of the Danian, Ma-Da DS and
Da-1 DS, are Valvulineria patalaensis associated with
P.? antiqua, P.? cretae and Stomatorbina? binkhorsti
(samples Urb 1, Urb 2, Urb 3 and Urb 4; Fig. 11). It is
important to remark that Planorbulina? antiqua is located
in our material in the stratigraphic interval belonging to the
SBZ 1 (Urbasa Pass section, samples Urb 1, Urb 2 and Urb
3; Fig. 11) and in the lower part of the SBZ 2 (Urko section,
sample Ur 4; Fig. 15).
In sum, Laffitteina bibensis is an exclusively Cretaceous
species while the association of Bangiana hanseni,
Miscellanites globularis and Ornatononion moorkensii
does not allow anymore to propose the former species as
exclusive of SBZ 1. This zone is thus characterized by the
absence of the true larger foraminifera. Even the small
foraminifera, such as P.? antiqua, Valvulineria patalaensis
and S.? binkhorsti are not exclusive of this zone.
Remarks on SBZ 2. Serra Kiel et al. (1998)
characterized biozone SBZ 2 based on the assemblage
formed by Miscellanea globularis, Ornatononion
minutus, Paralockhartia eos and Lockhartia akbari.
However, according to Hottinger (2009) Miscellanea
globularis has a biostratigraphic range encompassing SBZ
2 and SBZ 3, a fact that rules it out as a marker for SBZ 2.
In turn, the species identified as Ornatononion minutus
by Serra-Kiel et al. (1998) and combined as Miscellanea
minuta by Rahaghi (1983; pl. 43, figs. 6-11) would be
restricted to SBZ 3 according to Hottinger (2009), excluding
it from SBZ 2. Moreover, Paralockhartia eos would have
a biostratigraphic range restricted to SBZ 2 according to
Hottinger (2014). Other larger foraminifera found in the
material studied and belonging to SBZ 2 shed new light on
the assemblage characteristic of this biozone:
Elazigina dienii (=Plumokathina dienii), attributed
exclusively to SBZ 2 by Hottinger (2014), has been found
associated with Miscellanites globularis in Garralda
section (sample G 2; Fig. 7), with Haymanella elongata,
Rotorbinella conica, Daviesina praegarumnensis and

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Paralockhartia eos in Lizarraga section (Fig. 10), with
Miscellanites globularis, Stomatorbina? binkhorsti,
Redmondina henningtoni and Valvulineria patalaensis
in Aixola-Ermua section (Fig. 14) and with M. globularis
and Ornatononion moorkensii in Urko section (Fig. 15).
These results confirm the exclusive attribution of E. dienii
to SBZ 2.
Also, Ornatononion moorkensii is confirmed to be
restricted to the SBZ 2 as indicated by Hottinger (2014).
This species has been found associated with Haymanella
elongata, Scarificatina reinholdi and Planorbulina?
cretae in Lizarraga section (Fig. 10), with Rotorbinella
hensoni, Rotorbinella skourensis, Planorbulina? cretae,
Pyrenerotalia depressa, Kathina aquitanica, Elazigina
dienii,
Stomatorbina?
binkhorsti, Miscellanites
globularis, Cuvillierina sireli, Paralockhartia eos and
Scarificatina reinholdi in Urko section (Fig. 15).
Paralockhartia eos, attributed to the SBZ 2 by
Hottinger (2014), is also confirmed as a marker of SBZ 2 as
it has been found associated with Haymanella elongata,
Rotorbinella conica and Elazigina dienii in the Lizarraga
section (Fig. 10) and in the Urko section (Fig. 15). Also
Rotospirella conica belongs exclusively to the SBZ 2 and
has been found associated with Haymanella elongata,
Elazigina dienii, Daviesina praegarumnensis and
Paralockhartia eos in the Lizarraga section (Fig. 10).
The same situation is that of Pyrenerotalia depressa,
which has been found exclusively in rocks attributed to the
SBZ 2 in Urko section (Fig. 15) along with Rotorbinella
hensoni, Rotorbinella skourensis and Ornatononion
moorkensii.
On the other hand, some other associated species can
no longer be considered as characteristic of SBZ2. Thus,
Daviesina praegarumnensis extends from SBZ 2 to SBZ
3 according to (Hottinger, 2014), and it has been found
indeed in beds attributed to the SBZ 2 in Lizarraga section
(sample 3; Fig. 10) while in Campo section (samples Cm 3
and Cm 6; Fig. 5) it reaches the SBZ 3.
Other taxa that we found within the interval
characterized as SBZ 2 are Kayseriella decastroi
associated with Haymanella paleocenica, Haymanella
elongata, Scarificatina
reinholdi, Stomatorbina?
binkhorsti and Miscellanites globularis in Urbasa
Pass section (Fig. 11), associated with Miscellanites
globularis (Fig. 10) and Valvulineria patalaensis along
with Scarificatina reinholdi in Lizarraga section (Fig. 10)
and with Rotorbinella hensoni, Rotorbinella skourensis
and Scarificatina reinholdi in the Monte Urko section
(Fig. 15). Finally, Goesella sp. is present in the Urbasa Pass
section (Fig. 11) associated with Haymanella paleocenica,
H. elongata and Bangiana hanseni.

58

J. Serra-Kiel et al.

In sum, the new assemblage of species proposed
as markers for SBZ 2 is formed by Elazigina dienii,
Haymanella
elongata, Haymanella
paleocenica,
Kayseriella decastroi, Ornatononion moorkensii,
Pyrenerotalia depressa, Rotorbinella conica and
Paralockhartia eos.
Remarks on SBZ 3. Serra-Kiel et al. (1998)
characterized the SBZ 3 with the assemblage formed
by Glomalveolina primaeva, Periloculina slovenica,
Coskinon rajkae, Fallotella alavensis, Cribrobulimina
carniolica, Vania anatolica, Miscellanea yvettae,
Pseudomiscellanea primitiva, Ranikothalia bermudezi,
“Operculina” heberti and Discocyclina seunesi. The
data obtained here indicate that this assemblage shoul be
modified in order to characterize properly the SBZ 3 as
follows.
•	
Glomalveolina primaeva, was found in the
sections of Serraduy (Fig. 4), Campo (Fig. 5), Tena (Fig. 6),
Andia (Fig. 9), Lizarraga (Fig. 10), Urbasa Pass (Fig. 11),
Leortza (Fig. 12) and Aixola-Ermua (Fig. 14).
•	
Periloculina slovenica is present in the sections
of Campo (Fig. 5), Tena (Fig. 6), Andia (Fig. 9), Lizarraga
(Fig. 10) and Leortza (Fig. 12).
•	
Coskinon rajkae is present in Campo section
(Fig. 5), Tena section (Fig. 6), Andia section (Fig. 9),
Lizarraga section (Fig. 10) and Leortza section (Fig. 12).
•	
Fallotella alavensis is present in the sections of
Campo (Fig. 5), Tena (Fig. 6), Leortza (Fig. 12) and AixolaErmua (Fig. 14).
•	
Cribrobulimina carniolica occurs in the sections
of Serraduy (Fig. 4), Campo (Fig. 5), Tena (Fig. 6) and
Leortza (Fig. 12).
•	
Vania anatolica occurs in Campo section (Fig.
5), Tena section (Fig. 6) and Leortza section (Fig. 12).
•	
Miscellanea yvettae is present in Campo section
(Fig. 5), Garralda section (Fig. 7) and Leortza section
(Fig. 12).
•	
Miscellanea juliettae is present in Urbasa Pass
(Fig. 11) and the Leortza section (Fig. 12).
•	
Miscellanites primitivus (=Pseudomiscellanea
primitiva), was found in the sections of Garralda (Fig.
7), Andia (Fig. 9), Lizarraga (Fig. 10) and Urbasa Pass
(Fig. 11).
•	
Miscellanites minimus is present in Garralda
section (Fig. 7).
•	“Operculina” heberti is a relatively common
taxon found in Tena section (Fig. 6), Urrobi section (Fig. 8),
Andia section (Fig. 9), Lizarraga section (Fig. 10), Urbasa
Pass (Fig. 11), Leortza section (Fig. 12), Aixola-Ermua
section (Fig. 14) and the Bouzin outcrop.
•	
Discocyclina seunesi is present in the Tena
section (Fig. 6), the Urrobi section (Fig. 8), the Andia
section (Fig. 9), the Lizarraga section (Fig. 10), the

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Paleocene LBF from the Pyrenean Basin

Urbasa Pass section (Fig. 11) and the Leortza section
(Fig. 12).
•	
Ranikothalia soldadensis occurs in Urbasa Pass
(Fig. 11) and the Ruisseau de la Mède outcrop.
Other characteristic species located within the SBZ 3
interval are:
•	
Idalina sinjarica, found in the sections of
Serraduy (Fig. 4), Campo (Fig. 5), Tena (Fig. 6), Lizarraga
(Fig. 10) and Leortza (Fig. 12).
•	
Pseudolacazina donatae, present in Campo
(Fig. 5), Tena (Fig. 6) and Leortza (Fig. 12).
•	
Cuvillierina sireli, in Garralda (Fig. 7), Urbasa
pass (Fig. 11) and Leortza (Fig. 12).
•	
Scarificatina reinholdi, only present in Lizarraga
(Fig. 10).
•	
Redmondina henningtoni, in Campo (Fig.
5), Tena (Fig. 6), Leortza (Fig. 12) and Aixola-Ermua
(Fig. 14).
•	
Elazigina lenticula, found in Aixola-Ermua
(Fig. 14), Andia (Fig. 9) and Lizarraga pass (Fig. 10).
•	
Cincoriola ovoidea, present in Campo (Fig. 5).
•	
Kathina aquitanica, present in the sections of
Serraduy (Fig. 4), Campo (Fig. 5), Garralda (Fig. 7) and
Andia (Fig. 9).
•	
Kathina pernavuti was only found in the Campo
section (Fig. 5).
•	
Thalmannita madrugaensis, is present in Tena
section (Fig. 6), Urrobi section (Fig. 8), Urbasa Pass
section (Fig. 11), Leortza section (Fig. 12) and AixolaErmua (Fig. 14).
•	
Daviesina praegarumnensis, occurs in Campo
section (Fig. 5).
•	
Valvulineria bacetai n. sp., is present in Campo
(Fig. 5), Lizarraga pass (Fig. 10) and Urbasa Pass (Fig. 11).
It should be noted that, according to Hottinger (2014), the
biostratigraphic ranges of Kathina lenticula, Thalmannita
madrugaensis and Redmondina henningtoni are from
SBZ 3 to SBZ8, from SBZ 3 to SBZ 4 and from SBZ 3 to
SBZ 6 respectively.
In sum, the new assemblage proposed to characterize
SBZ 3 is formed by Glomalveolina primaeva, Periloculina
slovenica, Vania anatolica, Coskinon rajkae, Fallotella
alavensis, Cribrobulimina carniolica, Miscellanea
yvettae, Miscellanea juliettae, Miscellanites primitivus,
Miscellanites minutus, Ranikothalia soldadensis,
“Operculina” heberti and Discocyclina seunesi.
Remarks on SBZ 4. Serra-Kiel et al. (1998)
characterized SBZ 4 biozone with the assemblage formed
by Glomalveolina levis, Hottingerina lukasi, Miscellanea
meandrina, Daviesina garumnensis, Dictyokathina
simplex, Nummulites catari, Assilina azilensis and

59

R. hensoni

S. iranicus

paleocenica

“O.”

Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

FIGURE 28.Paleocene larger foraminifera distribution. The range shown with a red line is in accordance with the indications of previous authors.

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Assilina yvettae. It should be noted that, after Hottinger
(2009), Miscellanea meandrina has been combined as
Miscellanites meandrinus. The proposal of combining
this species within genus Diyadinella by Sirel (2018) is
not well documented from our point of view.
According to the stratigraphic distribution of larger
foraminifera in this work, the SBZ 3 biozone is characterized
by the presence of following markers:
•	
Glomalveolina levis, in Serraduy section (Fig. 4),
in Campo section (Fig. 5), in Tena section (Fig. 6), in Korres
section (Fig. 13).
•	
Alveolina korresensis, in Korres section (Fig. 13)
•	
Hottingerina lukasi, in Korres section (Fig. 13).
•	
Daviesina garumnensis, in Serraduy section (Fig.
4), in Campo section (Fig. 5), in Korres section (Fig. 13), in
Aixola-Ermua section (Fig. 14) and samples from Le Quillet
and Cérisols outcrops.
•	
Assilina yvettae, in Campo section (Fig. 5), in
Tena section (Fig. 6), in Urrobi section (Fig. 8), in in Korres
section (Fig. 13), in Aixola-Ermua section (Fig. 14) and
sample from Le Quillet outcrop.
•	
Assilina azilensis, in Campo section (Fig. 5), in
Tena section (Fig. 6), in Urrobi section (Fig. 8), in Urbasa
Pass section (Fig. 11), in Leortza section (Fig. 12), in
Aixola-Ermua section (Fig. 14) and samples from Cérisols
and Le Quillet outcrops.
•	
Nummulites catari, samples from Cérisols and
Le Quillet outcrops.
Other species located within the interval characterized
as SBZ 4 we found:
•	
Elazigina lenticula in Korres section (Fig. 13), in
Aixola-Ermua section (Fig. 14).
•	
Elazigina subsphaerica, in Aixola-Ermua
section (Fig. 14).
•	
Valvulineria bacetai n. sp., in Korres section
(Fig. 13).
•	
Redmondina henningtoni, in Korres section (Fig. 13).
•	
Sistanites iranicus, in Aixola-Ermua section
(Fig. 14).
In sum, the new assemblage proposed to characterize
SBZ 4 is formed by Glomalveolina levis, Alveolina
korresensis,
Hottingerina
lukasi,
Daviesina
garumnensis, Assilina yvettae, Assilina azilensis and
Nummulites catari.
The study of calcareous nannoplankton (NP and CNP
zones) and planktonic foraminifera (P zones) allows for a
correlation of these zones with the SBZ.
The SBZ 1 biozone can be correlated with calcareous
plankton in the Aixola-Ermua and Monte Urko sections.

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

In Aixola-Ermua it encompasses the interval pertaining
to the lower part of P1 zone (P1a+b subzones; Fig. 14),
while in Monte Urko section, the SBZ 1 is correlated with
the lower-middle part of calcareous nannoplankton NP2 or
CNP2-partim CNP3 zones (Fig. 15). These data indicate
that the top of the SBZ 1 is located within the upper part
of NP2 zone, whereas its base at an indeterminate point
within this zone. In the Aixola-Ermua section, SBZ 1 is
correlated with the lower part of planktonic foraminifera
zone P1 (P1a+P1b in Fig. 14).
The SBZ 2 biozone is correlated with calcareous
nannofossils in the Urko section, in the interval comprised
between the upper part of NP2 and the upper part of
NP 4, or the lower part of CNP3 and the beginning of
CNP7 (Fig. 15), and in part with P1c to P3 zones of the
planktonic foraminifera in the Aixola-Ermua section (Fig.
14). Biostratigraphic data from the platform margin series
studied by Baceta et al. (2005) document a correlation
of the SBZ 2 with calcareous nannoplancton biozones
spanning from the middle part of NP3 to the uppermost
of the NP4. The discrepancy in the position of the base
of the SBZ 2 as to that defined at the Monte Urko section
most likely derives from the absence of shallow benthic
foraminifera in the Da1 DS deposits exposed at Lizarraga
pass section.
The SBZ 3 biozone is correlated with the biostratigraphic
interval between the lowermost NP 5 and the topmost NP
8 zones (partim CNP7-partim CNP10) of calcareous
nannoplankton in the Andia and Aixola-Ermua sections
(Figs. 9; 14) and with the P4 planktonic foraminiferal zone
in the late section. Finally, SBZ 4 biozone is correlated
with partim NP9 or partim CNP11 zones of calcareous
nannoplankton in the Campo and Aixola-Ermua sections
(Figs. 5; 14).
The Paleocene-Eocene boundary is represented by the
geochemical signal of a CIE according to Orue-Etxebarria
et al. (2001) and Pujalte et al. (2009b). This excursion is
located in the Campo, Urrobi and Aixola-Ermua sections
(Figs. 5; 8; 14) according to Schmitz and Pujalte (2003),
Pujalte et al. (2003) and Pujalte et al. (2009a, b).
The biostratigraphic range of the species studied in this
work is represented in Figure 28, in accordance with the
International Chronostratigraphic Chart 2017 (Cohen et al.
2013; Vandenberghe et al. 2012).

CONCLUSIONS
Detailed taxonomic study of the Paleocene larger
foraminifer from the Pyrenean basin has allowed us to
identify sixty taxa, including two new species of larger

61

Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

foraminifera: Alveolina korresensis n. sp. and Valvulineria
bacetai n. sp.
The present study has allowed us to redefine and
recalibrate the SBZs of the Paleocene according to new
correlation data with calcareous nannoplancton and
planktnic foraminifera, as follows (Fig. 29):
•	
SBZ 1. According to the results of this study,
Laffitteina bibensis and Bangiana hanseni can be
definitively ruled out as markers for SBZ 1, contrary to the
indications in Serra-Kiel et al., (1998). The SBZ 1 has,
therefore, no exclusive markers and is now redefined to be
characterized by an assemblage composed of Valvulineria
patalaensis, Stomatorbina? binkhorsti, Planorbulina?
antiqua and Bangiana hanseni. This assemblage
encompasses the interval between the K/Pg boundary and
the upper part of the Danian desposits (Da-2 DS). SBZ 1
is constrained by the chronostratigraphic interval between

the K/Pg boundary and an intermediate point between the
CNP2-CNP3 and the NP2-NP3 boundaries, corresponding
to the first appearance of the markers of the SBZ 2.
•	
SBZ 2 is characterized by the assemblage
of Haymanella elongata, Haymanella paleocenica,
Kayseriella decastroi, Rotospirella conica, Pyrenerotalia
depressa Elazigina dienii, Ornatononion moorkensii and
Paralockhartia eos. This zone is reasigned in age to the
interval from topmost NP2 to uppermost NP4 calcareous
nannofossil zones (CNP3 to lowermost CNP7, Fig. 29).
•	
SBZ 3 is characterized by the assemblage of
Glomalveolina primaeva, Periloculina slovenica,
Vania anatolica, Coskinon rajkae, Fallotella alavensis,
Cribrobulimina carniolica, Miscellanea yvettae,
Miscellanea
juliettae,
Miscellanites
primitivus,
Miscellanites minutus, Ranikothalia soldadensis,
“Operculina” heberti and Discocyclina seunesi. This

Planktic Foraminifera

“O.”

P1

c

a+b
P0+Pα

FIGURE 29. Correlation between the Geological Time Scale, magnetostratigraphy, planktic foraminifera zones (P), calcareous nannoplankton zones
(NP, CNP), shallow benthic zones (SBZs) according to Serra-Kiel et al. (1998), the SBZs in this work, Paleocene main larger foraminifera markers
and the depositional sequences (DS).

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

62

Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

zone encompasses from lower NP5 to topmost NP8
calcareous nannofossil zones (CNP7 to CNP10, Fig. 29).
•	
SBZ 4 is characterized by the assemblage of
Glomalveolina levis, Alveolina korresensis, Hottingerina
lukasi, Daviesina garumnensis, Assilina yvettae,
Assilina azilensis and Nummulites catari. In age this
zone encompasses the NP9 calcareous nannofossil zone
(most of CNP11, Fig. 29)
The correlation between larger foraminifera and
calcareous nannoplankton in Campo, Andia, Lizarraga,
Aixola-Ermua and Urko sections has allowed the
calibration of the SBZs with the Geologic Time Scale
produced in accordance with the 2017 International
Chronostratigraphic Chart (Cohen et al., 2013;
Vandenberghe et al., 2012) (Fig. 29).
The main modifications of the biostratigraphic range of
the Paleocene SBZ with the Serra-Kiel et al. (1998) are as
follows (Fig. 29):
•	 Reduction of the SBZ 1 range from 4.06m.y. to
1.09m.y.
•	 Extension of the SBZ 2 range from 1.19m.y. to
3.34m.y.
•	
Extension of the SBZ 3 range from 2+
•	
The SBZ 3 covers from the base of the Selandian
to the middle part of the Thanetian. The SBZ 4 is late
Thanetian.
Furthermore, the combined study of calcareous
nannoplankton and larger foraminifera improve the
correlation between the DS as defined by Baceta et
al. (2004, 2005, 2011) and Schmitz et al. (2011), and
the Geological Time Scale (Fig. 2). Thus the ages of
these depositional sequences is as follows: Ma-Da DS
is Maastrichtian−earliest Danian in age; Da-1 DS is
early Danian; Da-2 DS is late Danian; Se/Th-1 DS is
Selandian/early−middle Thanetian and Th-2 D is late
Thanetian in age.
ACKNOWLEDGMENTS
We acknowledge the financial support for this
contribution from the Spanish Ministry of Economy
and Competitiveness projects CGL 2015-69805-P and
CGL2015-65404-R. This is also a contribution to the
Research Group of the Basque University System IT-93016. This study was also conducted in the framework of the
research and collections projects of the Museu de Ciències
Naturals de Barcelona (MCNB). Sincere thanks to the
reviewers Johannes Pignatti and Josep Tosquella for their
constructive comments and suggestions.

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

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Notebooks on Geology, 14(4), 41-68.
Vicedo, V Robles-Salcedo, R., Serra-Kiel, J., Hidalgo, C., Razin,
.,
Ph., Grélaud, C., 2019. Biostratigraphy and evolution of the
larger rotaliid foraminifera in the Cretaceous-Paleogene
transition of the southern Oman Mountains. Papers in
Paleontology, 1-43.
Villatte, J., 1962. Etude stratigraphique et paléontologique du
Montien des Petites Pyrénées et du Plantaurel. Toulouse,
Edouard Privat Maitre-Imprimeur, 1-331.
Villatte, J., 1969. Découverte d’une espèce du genre
Nummulitoides Abrard 1955 dans la zone a “Operculina”
heberti-Discocyclina seunesi du Thanétien des PetitesPyrénées. Compte Rendu Sommaire Séances Société
géoligique de France, 9(1968), 299-300.
Wan, X., 1991. Paleocene larger foraminifera from Southern Tibet.
Revista Española de Micropaleontología, 23, 7-28.
Wright, V ., Burchette, T.P 1996. Shallow-water carbonate
.P
.,
environments. In: Reading, H.G. (ed.). Sedimentary
environments. 3rd Edition Blackwell Scientific, 325-394.
Manuscript received June 2019;
revision accepted April 2020;
published Online June 2020.

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

69

Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

APPENDIX I

TABLE I. Morphometrically parameters measured in the equatorial sections of megalospheric forms of the species Nummulites catari Tosquella and
Serra-Kiel, 1998

Whorl (nº)

1

3

0.41-0.53

0.56-1.11

0.81-1.52

ago-16

Radius (mm)

2

37-48

79-103

Septa (nº)

TABLE II. Morphometrically parameters measured in the equatorial sections of megalospheric forms of the species Assilina yvettae (schaub, 1981)

Whorl (nº)

1

3

0.38

0.75

1.20

10

Radius (mm)

2

29

53

Septa (nº)

TABLE III. Morphometrically parameters measured in the equatorial sections of megalospheric forms of the species Assilina azilensis (tambareau,
1966)

Microspheric generation
Whorl (nº)

1

2

3

4

5

0.1

0.3

0.67

1.45

2.94

Septa (nº)
8
Megalospheric generation

24

44

85

155

1

2

3

-

-

0.54-0.61

1.08-1.23

1.77-1.99

-

-

07-nov

23-32

48-59

-

-

Radius (mm)

Whorl (nº)
Radius (mm)
Septa (nº)

TABLE IV. Morphometrically parameters measured in the equatorial sections of megalospheric forms of the species Ranikothalia soldadensis (vaughan
and cole, 1941)

Microspheric generation
Whorl (nº)

1

2

3

4

0.47

1.16

2.99

3.34

Septa (nº)
11
Megalospheric generation

29

52

87

1

2

3

-

0.50-0.52

1.14-1.19

1.6-1.76

-

04-may

20

46-49

-

Radius (mm)

Whorl (nº)
Radius (mm)
Septa (nº)

Geologica Acta, 18.8, 1-69, I-III (2020)
DOI: 10.1344/GeologicaActayear2020.18.8

I

Paleocene LBF from the Pyrenean Basin

J. Serra-Kiel et al.

TABLE V. Samples.

Sample

Repository

Abau 1
Instituto Geológico y Minero de España (IGME)
Ain 30
Serra‐Kiel collection.Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ain 5
Instituto Geológico y Minero de España (IGME)
Aix 12
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Aix 14
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Aix 16
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Aix 19
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Aix 5
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Aix 9
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
An 1
Instituto Geológico y Minero de España (IGME)
An 10
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
An 8
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
An 9
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Beo 2
Instituto Geológico y Minero de España (IGME)
Beo 4
Instituto Geológico y Minero de España (IGME)
Beo 5
Instituto Geológico y Minero de España (IGME)
Bin 3
Instituto Geológico y Minero de España (IGME)
Bouzin
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Bur 3
Instituto Geológico y Minero de España (IGME)
Bur 7
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Campeche (Mexico)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cérisols  (Petites Pyrénées)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cm 2
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cm 3
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cm 4
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cm 5
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cm 6
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cm 7
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Cm 8
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Coll 3
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Dor M'Said
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Er 1
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Er 5
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Eu 2
Instituto Geológico y Minero de España (IGME)
G 3
Instituto Geológico y Minero de España (IGME)
K 1
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
K 2
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
L 12
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 13
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 14
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 15
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 17
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 18
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 19
L 20
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
L 21
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 22
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 24
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 25
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 26
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 27
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 3
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 4
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 5
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 2
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 5
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 7
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 8
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Le Quillet  (Petites Pyrénées)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Geologica Acta, 18.8, 1-69, I-III (2020)
Narp (Petites Pyrénées)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
D O I : 1 0 . 1 3 4 4 / G e o l o g i c a ADepartment of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
ctayear2020.18.8
Orio 3
RM‐Le 12
Instituto Geológico y Minero de España (IGME)
Ruisseau de la Mède  (Petites 
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Pyrénées)

II

L 21
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 22
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 24 r r a - K i e l e t a l .
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Paleocene LBF from the Pyrenean Basin
J. Se
L 25
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 26
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 27
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 3
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
TABLE V. Continued.
L 4
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 5
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 2
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Sample
Repository
Le 5
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Abau 1
Instituto Geológico y Minero de España (IGME)
Le 7
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Ain 30
Serra‐Kiel collection.Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Le 8
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ain 5
Instituto Geológico y Minero de España (IGME)
Le Quillet  (Petites Pyrénées)
Aix 12
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Narp (Petites Pyrénées)
Aix 14
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Orio 3
Aix 16
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
RM‐Le 12
Instituto Geológico y Minero de España (IGME)
Aix 19
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ruisseau de la Mède  (Petites  Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Aix 5
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Pyrénées)
Aix 9
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Salt Range (Pakistan)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
An 1
Instituto Geológico y Minero de España (IGME)
Salv 4
Instituto Geológico y Minero de España (IGME)
An 10
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Salv 5
Instituto Geológico y Minero de España (IGME)
An 8
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Tap 10
Instituto Geológico y Minero de España (IGME)
An 9
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Tap 15
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Beo 2
Instituto Geológico y Minero de España (IGME)
Te 1
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Beo 4
Instituto Geológico y Minero de España (IGME)
Te 4
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Beo 5
Instituto Geológico y Minero de España (IGME)
Ur 10
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Bin 3
Instituto Geológico y Minero de España (IGME)
Ur 15
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Bouzin
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ur 17
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Bur 3
Instituto Geológico y Minero de España (IGME)
Ur 18
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Bur 7
Ur 3
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Campeche (Mexico)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ur 4
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Cérisols  (Petites Pyrénées)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ur 6
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Cm 2
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ur 8
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Cm 3
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Ur 9
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Cm 4
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 1
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Cm 5
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Urb 12
Cm 6
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 13
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Cm 7
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 15
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Cm 8
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 18
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Coll 3
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 19
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Dor M'Said
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 20
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Er 1
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 25
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Er 5
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 26
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Eu 2
Instituto Geológico y Minero de España (IGME)
Urb 27
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
G 3
Instituto Geológico y Minero de España (IGME)
Urb 31
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
K 1
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urb 4
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
K 2
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Urrob 1
Instituto Geológico y Minero de España (IGME)
L 12
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Urrob 3
Instituto Geológico y Minero de España (IGME)
L 13
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 14
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 15
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 17
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 18
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 19
L 20
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
L 21
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 22
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 24
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 25
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 26
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 27
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 3
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 4
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
L 5
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 2
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 5
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 7
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
Le 8
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Le Quillet  (Petites Pyrénées)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
III
Geologica Acta, 18.8, 1-69, I-III (2020)
Narp (Petites Pyrénées)
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
DOI: 10.1344/GeologicaActayear2020.18.8
Orio 3
Department of Stratigraphy and Paleontology, University of the Basque Country (UPV/EHU) 
RM‐Le 12
Instituto Geológico y Minero de España (IGME)
Ruisseau de la Mède  (Petites 
Serra‐Kiel collection. Department of Earth and Ocean Dynamics, University of Barcelona (UB)  
Pyrénées)