The bathyal larger lituolid Neonavarella n. gen. (Foraminifera) from the Thanetian Scaglia Rossa Formation of northeastern Italy

Abstract

Larger agglutinated foraminifera resembling the Cretaceous genus Navarella Ciry and Rat 1951 were recently recovered
in Thanetian hemipelagites from the Belluno Basin, northeastern Italy. These lituoids first appear in the basal Thanetian (uppermost
calcareous nannofossils Zone CNP 8) and become common in the >500 μm washed residue from the uppermost Thanetian. They
abruptly disappear at the Paleocene/Eocene boundary, in coincidence with the extinction of Paleocene small benthic foraminifera (the
benthic foraminiferal extinction event - BEE).
In order to document the internal chamber arrangement and the agglutinated wall microstructure of the Thanetian lituolids and to
compare them with similar individuals recovered from the Upper Cretaceous and Danian strata of the same section, the collected specimens
were sectioned and analyzed using a Scanning Electron Microscope (SEM), equipped with an energy-dispersive X-ray spectrometer
(EDX). Our results show a typical bi-layered wall microstructure in the Thanetian specimens, whereas the older Maastrichtian and
Danian specimens, occurring in the same section, display a single, thicker agglutinated wall. The taxonomy of the Italian lituolids is discussed
and compared with similar taxa known from the literature.
We describe the Thanetian lituolids as the new genus Neonavarella, which shows an apparently identical external morphology to
mono-layered Maastrichtian–Danian specimens but differs in the microstructure of the agglutinated test wall that is bi-layered. The
finding of new and well-preserved material from the Paleocene Scaglia Rossa beds of Italy helps shed light on the taxonomy of the still
poorly known deep-water larger lituolids.

Full text

The bathyal larger lituolid Neonavarella n. gen.
(Foraminifera) from the Thanetian Scaglia Rossa Formation
of northeastern Italy
Luca Giusberti1, Michael A. Kaminski2and Nicoletta Mancin3
1Department of Geosciences, University of Padova, via Gradenigo 6, I-35131 Padova, Italy
email: luca.giusberti@unipd.it
2Geosciences Department, King Fahd University of Petroleum & Minerals,
PO Box 701 KFUPM, Dhahran, 31261, Saudi Arabia
email: kaminski@kfupm.edu.sa
3Department of Earth and Environment Sciences, University of Pavia, via Ferrata 9, 27100 Pavia, Italy
corresponding author email: nicoletta.mancin@unipv.it
ABSTRACT: Larger agglutinated foraminifera resembling the Cretaceous genus Navarella Ciry and Rat 1951 were recently recov-
ered in Thanetian hemipelagites from the Belluno Basin, northeastern Italy. These lituoids first appear in the basal Thanetian (upper-
most calcareous nannofossils Zone CNP 8) and become common in the >500 μm washed residue from the uppermost Thanetian. They
abruptly disappear at the Paleocene/Eocene boundary, in coincidence with the extinction of Paleocene small benthic foraminifera (the
benthic foraminiferal extinction event - BEE).
In order to document the internal chamber arrangement and the agglutinated wall microstructure of the Thanetian lituolids and to
compare them with similar individuals recovered from the Upper Cretaceous and Danian strata of the same section, the collected speci-
mens were sectioned and analyzed using a Scanning Electron Microscope (SEM), equipped with an energy-dispersive X-ray spectrom-
eter (EDX). Our results show a typical bi-layered wall microstructure in the Thanetian specimens, whereas the older Maastrichtian and
Danian specimens, occurring in the same section, display a single, thicker agglutinated wall. The taxonomy of the Italian lituolids is dis-
cussed and compared with similar taxa known from the literature.
We describe the Thanetian lituolids as the new genus Neonavarella, which shows an apparently identical external morphology to
mono-layered Maastrichtian–Danian specimens but differs in the microstructure of the agglutinated test wall that is bi-layered. The
finding of new and well-preserved material from the Paleocene Scaglia Rossa beds of Italy helps shed light on the taxonomy of the still
poorly known deep-water larger lituolids.
Key words: Larger agglutinated foraminifera, lituolids, Systematics; New genus, New species, Upper Cretaceous, Paleocene, north-
eastern Italy.
INTRODUCTION
Since the 19th century, large-sized lituolids have been widely re-
ported in Upper Cretaceous turbiditic and hemipelagic succes-
sions of the Tethyan domain (e.g., Ciry and Rat 1951; Maync
1952; 1954; Cati 1960; Sampò 1972; Stacher 1980; Pandey
1981; Riegraf 1998; Radoi¹iºet al. 2010). Such agglutinated
foraminifera were generally ascribed to different species of the
genera Lituola Lamarck 1804, Navarella Ciry and Rat 1951 and
Recurvoides Earland 1934.
In the course of a study of the Paleocene/Eocene Thermal Maxi-
mum as recorded in a “Scaglia-type” bathyal succession of the
Southern Alps of northeastern Italy (Forada section), several hun-
dred specimens of larger agglutinated lituolids were detected for
the first time in Thanetian hemipelagic rocks (Giusberti et al.
2016). After intensive research performed during this study, simi-
lar specimens were also sporadically observed in Danian
hemipelagites of the same section, though with very rare abun-
dance. The Paleocene specimens, generally larger than 500 μm,
are virtually indistinguishable from Upper Cretaceous specimens
recovered elsewhere and described in the literature as Navarella.
Navarella is a larger lituolid belonging to the family Ammo-
baculinidae Saidova 1981 and attaining a maximum length of 5
mm, with an initially streptospirally enrolled test, later uncoiled,
and with an aperture varying during ontogeny from slit-like to
cribrate. The genus was originally described from the Maast-
richtian flysch of the Navarra region in the Spanish Pyrenees,
and then reported elsewhere in Europe from Campanian to
Maastrichtian rocks. The validity of this taxon and its range dis-
tribution is, however, controversial and strongly debated
(Riegraf 1998).
In this study, we use high definition SEM EDX images of entire
and sectioned specimens, and study the external and internal
features of the Paleocene lituolids in order to investigate both
their test morphology and microstructure of the agglutinated
wall. The main purpose is to compare the recorded Paleocene
specimens with similar Italian and foreign Cretaceous taxa in
order to verify whether or not the newly discovered aggluti-
nated foraminifera are ascribable to Navarella, thereby extend-
ing the known stratigraphical range of navarellids to the
Paleocene.
Micropaleontology, vol. 64, nos. 5-6, text-figure 1, table 1, plates 1–7, supplemental material online, pp. 417–434, 2018 417

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Luca Giusberti et al.: The bathyal larger lituolid Neonavarella n. gen. from the Thanetian Scaglia Rossa Formation, northeastern Italy
TEXT-FIGURE 1
A) location of the Forada section; B) stratigraphic log of the Forada section with the range of Thanetian Neonavarella n. gen. plotted against the isotope
curve of d13C. Red asterisks mark samples from Danian and upper Maastrichtian records where sporadic specimensof navarellid lituolids were recov-
ered; C) detail of the Thanetian–Ypresian portion of the Forada section plotted against the absolute abundance of Neonavarella n. gen. (number of speci-
mens per gram of dry sediment). CNP and CNE calcareous nannofossil zones of text-fig. 1B and 1C according to Agnini et al. (2014)

419
Micropaleontology, vol. 64, nos. 5–6, 2018
TABLE 1
Diagnostic features (external morphology, chamber arrangement, aperture and test wall microstructure) of larger lituolids described in the literature from Upper Cretaceous records of Europe, Italy and
in the present work from the Maastrichtian-Thanetian record of the Forada section (Belluno) and Monte San Lorenzo Quarry section (Maniago).

MATERIALS AND METHODS
Forada section
The investigated section (46.036083°N, 12.063975°E) crops
out along the Forada Creek, roughly 2 km east of the Lentiai
village in the Venetian Prealps (northeastern Italy), and consists
of ~62 m of pink-reddish Scaglia Rossa limestones and marly
limestones, locally rhythmically organized (text-fig. 1). It re-
cords a continuous stratigraphic interval from the uppermost
Cretaceous to the lower Eocene (upper Maastrichtian–lower
Ypresian nannofossil Zone CNE2 of Agnini et al. 2014) and is
virtually unaffected by structural complications (Fornaciari et
al. 2007; Giusberti et al. 2007).
At Forada, ichnofossils are well represented (e.g., Zoophycos,
Thalassinoides) as they generally occur in the upper portion of
the Scaglia Rossa Formation in the Belluno Basin (Miller
2000). The only macrofossils observed are rare irregular
echinoids and possibly an octocoral, all recovered in the Danian
beds (Giusberti et al. 2005; J. Stolarski, pers. comm.).
The lowermost Danian record, above the K/Pg boundary clay, is
represented by a subnodular/irregularly bedded red-brown lime-
stone and marly limestone, and has been object of a study aimed
to investigate the recovery of the calcareous plankton in the after-
math of the K/Pg boundary event (Fornaciari et al. 2007). The
rest of the Danian and Selandian succession is represented by red
carbonate and marly lithofacies, sometimes strongly bioturbated
by Zoophycos (“massive unit”); three thin biocalciturbiditic beds
(4.5 to 10 cm thick), with an erosive base, also characterize the
lower Danian portion of the section (text-fig. 1).
The Danian–Selandian “massive unit” is overlain by Thanetian
“scaly” reddish calcareous marls mottled by greenish “flames”
(“scaly unit”). At about -20 cm below the Paleocene/Eocene
boundary, the lithology shifts to greenish marls with Zoophycos
and Chondrites (intervals Pa I and Pa II of Giusberti et al.
2007). The entire Paleocene record, 40 m thick, is capped by
about 3.5 m thick greenish and reddish clay marl unit (CMU;
Giusberti et al. 2007), a lithological “anomaly” which marks the
so-called Paleocene/Eocene Thermal Maximum (PETM; e.g.,
McInerney and Wing 2011) in the study area (Giusberti et al.
2007; text-fig. 1). The CMU is followed by 4-5 m of marl-lime-
stones couplets, fading into 15 m of scaly, still strongly cyclical
reddish marls, Ypresian in age (text-fig. 1).
Based on benthic foraminiferal content, the upper Paleocene to
lower Eocene Scaglia Rossa interval has been probably depos-
ited at middle-lower bathyal paleodepth, likely between 1000
and 1500 m (Giusberti et al. 2016). The wealth of micropale-
ontological, sedimentological, and geochemical data so far col-
lected from the Forada section (e.g., Giusberti et al. 2007;
Agnini et al. 2007; Luciani et al. 2007; Tipple et al. 2011;
Giusberti et al. 2016) provides probably the most complete
paleoenvironmental reconstruction across the PETM in Europe
to date.
Agglutinated foraminifera
The larger lituolids here investigated were observed for the first
time by Giusberti et al. (2016) analyzing the >500 μm washed
fraction, originally used for the quantitative study of
microbenthic foraminifera across the Paleocene/Eocene Ther-
420
Luca Giusberti et al.: The bathyal larger lituolid Neonavarella n. gen. from the Thanetian Scaglia Rossa Formation, northeastern Italy
PLATE 1A
Intraspecific morphological variability of Paleocene lituolids from the Forada section.
Note that Thanetian and Danian specimens are morphologically identical. Images are in secondary electrons (SE).
1Thanetian juvenile specimen (MGP-PD 32076) with a
typical strongly involute coiling (sample BRI-6); a:
lateral view, b: profile. Note that the prime slit-like ap-
erture is still not manifest.
2 Thanetian juvenile specimen (MGP-PD 32076) with a
slit-like aperture (sample BRI-6); a: lateral view, b:
profile showing the slit-like aperture.
3 Thanetian quasi-adult specimen (MGP-PD 32077)
with a complex aperture characterized by a slightly
arched slit coupled with two small rounded pores
(sample BRI-6); a: lateral view, b: profile; c: detail of
the complex aperture.
4 Thanetian adult specimen (MGP-PD 32073) charac-
terized by a larger test with, a partly developed un-
coiled portion and a complex aperture formed by a
slightly arched slit coupled with some elongated pores
(sample BRI-29.5); a: lateral view, b: profile; c: detail
of the complex aperture.
5 Lateral view of a Thanetian adult specimen (MGP-PD
32073) with a typical large test characterized by a de-
veloped uncoiled portion (sample BRI-11);
6 Lateral view of an adult specimen (MGP-PD 32073)
with a very large test (exceeding 3 mm long and 1 mm
wide) characterized by a well developed uncoiled por-
tion (sample BRI-9).
7 Detail of the cribrate aperture of a Thanetian adult
specimen (MGP-PD 32081; sample BRI-9).
8 Lateral view of a Danian quasi-adult specimen
(MGP-PD 32074; RF-670, picked about 6.35 m above
the K/Pg boundary, Fig. 1).
9 Lateral view of a Danian juvenile specimen (MGP-PD
32074) very similar to the Thanetian specimen shown
in image 3a (sample RF-670).

Micropaleontology, vol. 64, nos. 5-6, 2018 421
Luca Giusberti, Michael A. Kaminski and Nicoletta Mancin Plate 1 A

422
PLATE 1B
Morphological details of upper Maastrichtian specimens of “Navarella” Ciry and Rat 1951 from the Forada section. To note that up-
per Cretaceous and Paleocene specimens seemingly possess very similar test morphologies. Images are in secondary electrons (SE).
1Adult specimen (MGP-PD 32089) characterized by a
large, ovoidal test (sample RF-24, just below the K/Pg
boundary; Fig. 1); a: lateral view, b: profile showing
the complex aperture formed by seven rounded or el-
liptical pores.
2Lateral view of an adult specimen (MGP-PD 32069)
showing a large test with the chambers initially spi-
rally arranged then becoming detached from the close
spire according to a rectilinear arrangement (sample
RF-24).
3 Lateral view of a small-size quasi-adult specimen
(MGP-PD 32070; sample RF-24).
PLATE 2
Sectioned specimens picked from the Thanetian record of the Forada section showing the internal chamber arrangement and
the microstructure of the agglutinated wall. In both juvenile and adult specimens the test wall results typically bi-layered, with
a thinner external portion made of coarse-grained mineral grains and a thicker, more homogeneous carbonatic portion, which
sometimes contains remains of planktonic foraminifera. Images of sectioned specimens are in back-scattered electrons (BSE).
1Juvenile specimen showing the chambers spirally ar-
ranged and the bi-layered microstructure of the agglu-
tinated wall (MGP-PD 32075; sample BRI-37); a:
lateral view, b: longitudinal section.
2Quasi-adult specimen (MGP-PD 32072) showing the
same bi-layered microstructure of the agglutinated
wall previously evidenced in the juvenile specimen
(sample BRI-9); a: lateral view, b: longitudinal sec-
tion; c: detail of the sectioned agglutinated wall with a
thicker internal carbonatic portion externally covered
by coarse-grained mineral granules.
3Adult specimen (MGP-PD 32071) showing the
chambers initially spirally arranged then becoming
detached from the close spire according to a rectilin-
ear arrangement and a more complex aperture (ar-
row). The microstructure of the agglutinated wall is
still bi-layered, with a thicker, internal carbonatic
portion and a thinner, external mineral pavement
(sample BRI-14.5); a: lateral view, b: longitudinal
section; c: detail of the sectioned agglutinated wall,
note in the carbonatic portion, a globigerinid test (ar-
row) filled by a white mineral (barite).
4Adult specimen (the same of Plate 1A, image 4a)
showing the uncoiled portion of the test and the typical
complex aperture (sample BRI-29.5); a: lateral view,
b: longitudinal section; c: detail of the thick bi-layered
wall (ca. 110 mm), d: thin section of the same speci-
men analysed at the polarized microscope showing
the bi-layered structure of the agglutinated wall
with the coarse-grained, external gains mostly
made of quartz (arrow).
5Adult specimen (the same of Plate 1A, image 6) show-
ing a very large test, with a particularly well devel-
oped uncoiled portion and the typical complex
aperture (sample BRI-9); a: lateral view, b: longitudi-
nal section showing the evolution of the aperture,
from a simple slit in the earlier chambers to complex
cribrate in the last chamber; c: detail of the thick
bi-layered wall (ca. 140 mm), d: thin section of the
same specimen analyzed at the polarized micro-
scope showing the bi-layered structure of the ag-
glutinated wall with the coarse-grained, external
gains mostly made of quartz.
Luca Giusberti, Michael A. Kaminski and Nicoletta Mancin Plate 1 B

Micropaleontology, vol. 64, nos. 5-6, 2018 423
Luca Giusberti, Michael A. Kaminski and Nicoletta Mancin Plate 2

mal Maximum. Very rare small juvenile lituolids were also ob-
served in the fraction >250 μm and <500 μm. The study of
Giusberti et al. (2016), based on 54 samples across an ~11 m
thick interval straddling the Paleocene/Eocene boundary (-467
to ca. +600 cm), revealed that these larger lituolids abruptly dis-
appear at the Paleocene/Eocene boundary, in coincidence with
the base of the CMU, where the benthic foraminiferal extinc-
tion (BEE) is recorded (Giusberti et al. 2016).
In order to verify the distribution of the investigated foram-
inifera throughout the entire section, 25 supplementary samples
were analyzed in search for the larger lituolids (text-fig. 1), in-
cluding those washed residues used for the study of foram-
inifera by Fornaciari et al. (2007) from the interval straddling
the Cretaceous/Paleogene (K/Pg) boundary. Furthermore, 8–9
new samples of a minimum weight of 200 g from the interval
richest in the investigated lituolids (ca. from -35.5 to -6 cm be-
low the P/E boundary) were treated for isolating a high number
of specimens for the present study.
Foraminifera were extracted from most of the indurate marls
and limestones using the “cold acetolysis” technique of Lirer
(2000) as described in Luciani et al. (2007); on the other hand
soft marly samples (the CMU, the greenish marl below the
CMU and some marly intercalations in the Thanetian portion of
the section) were prepared following standard procedures
(chemical disaggregation using a 10–30% solution of hydrogen
peroxide and subsequent washing through two sieves with
meshes of 500 and 63 mm). The absolute abundance of the
stated lituolids (N g-1: number of benthic foraminifera per gram
of bulk dried sediment) was calculated for the >500 μm fraction
in the interval 11 m thick straddling the P/E boundary (text-fig.
1).
SEM-EDS analysis
A total of about 240 lituolid specimens from the Paleocene re-
cord and 12 specimens from the uppermost Cretaceous portion
of the Forada section were isolated and identified, picked and
stored in micro-slides. Moreover, another three specimens were
isolated from the uppermost Thanetian Scaglia Rossa cropping
out at Monte San Lorenzo Quarry (Maniago, Pordenone prov-
ince, Friuli region; see Grandesso et al. 2008).
Following the methodology of Mancin et al. (2014), 61 repre-
sentative Paleocene specimens and 6 Maastrichtian specimens
from the Forada section (including one specimen from Monte
San Lorenzo Quarry) were mounted on stubs using carbon con-
ductive adhesive tapes and gold-coated for morphological anal-
yses (inBeam technique) by Scanning Electron Microscope
(SEM, Tescan FESEM, series Mira 3XMU) at the CISRiC-
Arvedi Laboratory (University of Pavia). Among the most rep-
resentative specimens previously analysed and photographed,
38 (respectively, 35 from the Paleocene and 3 from
Maastrichtian samples) were selected, then oriented and embed-
ded in epoxy resin, cut and polished with diamond pastes from
0.25 to 6 mm in grain-size and finally analysed using the SEM
equipped with an x-ray EDS. Back Scattered Electron (BSE)
images of sectioned specimens highlighted compositional simi-
larities (or dissimilarities) among agglutinated grains through
the wall thickness, on the basis of the mean atomic number of
each grain forming the agglutinated test. The elemental compo-
sition of the single grains forming the agglutinated wall was
provided through standardless spot microanalyses; moreover in
order to check the chemical-mineralogical variability of the ag-
glutinated grains within the sectioned test wall of each studied
specimen, two-dimensional x-ray maps of selected elements
424
Luca Giusberti et al.: The bathyal larger lituolid Neonavarella n. gen. from the Thanetian Scaglia Rossa Formation, northeastern Italy
PLATE 3
Sectioned specimens picked from Danian and upper Maastrichtian records of the Forada section showing the internal chamber
arrangement and the microstructure of the agglutinated wall. To note that in both Danian and Maastrichtian specimens the
microstructure of the test wall is evidently different with respect to Thanetian specimens and is characterized by a single, thicker,
homogeneous portion mostly made by carbonatic particles, sometimes containing tests of planktonic foraminifera.
1Danian juvenile specimen (the same of Plate 1A,
image 9); a: lateral view, b: longitudinal section (in
back scattered electrons-BSE) showing the cham-
bers spirally arranged and the simple slite-like ap-
erture (arrow), c: detail of the agglutinated wall
made of a single portion of fine-grained carbonatic
particles and tests of planktonic foraminifera.
2Danian quasi-adult specimen (the same of Plate 1A,
image 8); a: lateral view, b-c: longitudinal prepared as
thin section and photographed at the light polarized
microscope showing the chambers initially spirally
arranged, then becoming partly detached from the
close spire and the more complex aperture (arrows);
note that the agglutinated wall is homogeneous and
made by a single carbonatic portion (ca. 150 mm
thick).
3 Maastrichtian adult specimen (the same of Fig. Plate
1B, image 2); a: lateral view, b-c: longitudinal section
prepared as thin section and photographed at the light
polarized microscope (parallel and crossed nicols, b
and c respectively). Note the chambers that are at first
spirally arranged then becoming detached from the
close spire, note also the aperture that becomes more
complex during ontogeny (arrow). The agglutinated
wall is still homogeneous and made by a single
carbonatic portion.

Micropaleontology, vol. 64, nos. 5-6, 2018 425
Luca Giusberti, Michael A. Kaminski and Nicoletta Mancin Plate 3

were also collected. Each elemental map (silicon, calcium, alu-
minium, sodium, potassium and magnesium) was collected si-
multaneously, since the color intensity results proportional to
the element concentration in every pixel of the image (in black
areas the element is lacking) the comparison of the maps col-
lected in the same area of the sample provided an overview of
the mineralogical distribution in that area (Mancin et al. 2014).
Thin sections
Because most of the Cretaceous lituolids reported in the litera-
ture has been studied on thin sections, the most significant
Paleocene and Cretaceous specimens from the Forada section
were also prepared accordingly. Twenty-two specimens previ-
ously embedded in epoxy resin for EDS analysis (respectively,
15 from the Thanetian, 4 from the Danian and 3 from the
Maastrichtian) were cut and polished using standard hard rock
thin sectioning equipment and technique with minor modifica-
tions. Four 30 μm-thick thin sections pasted with epoxy onto a
glass slide were prepared for petrographic analysis using polar-
ized transmitted light optical microscopy. The studied speci-
mens (both isolated and sectioned) are housed in the
micropaleontological collections of the Museum of Geology
and Paleontology of Padova University (Italy).
RESULTS
Lituolid distribution pattern and foraminiferal assemblage
composition
Two sporadic occurrences of larger lituolids resembling to
Navarella have been observed in the Danian record of the
Forada section (at about + 6.35 and +16 m above the K/Pg
boundary; text-fig. 1); conversely no similar specimens occur in
the Selandian portion of the section. Rare specimens of larger
lituolids also occur in the Maastrichtian of the Forada section
exclusively at -24 cm below the K/Pg boundary (text-fig. 1).
The investigated lituolids appear consistently in the basal
Thanetian (calcareous nannofossils Zone CNP8 of Agnini et al.
2014) at about 21 m above the K/Pg boundary with an absolute
abundance of <0.05 specimen per gram, and become more
abundant at about -40 cm below the Paleocene/Eocene bound-
ary (up to ca. 0.1 specimen per gram). Moreover, these lituolid
specimens are a significant component of upper Paleocene ben-
thic foraminiferal assemblage in the >500 μm fraction, that
mostly consists of clavulinids, Cicicidoides, anomalinids and
trochamminids. This assemblage of larger-sized foraminifera,
including some characteristic Paleocene taxa, such as
Clavulinoides globulifera,Cibicidoides dayi and C. velasco-
ensis (CET, Cosmopolitan Extinction Taxa of Giusberti et al.
2016), abruptly disappear at about +39,5 m, in coincidence with
the onset of the negative carbon isotopic excursion of the
PETM and associated benthic foraminiferal extinction event
(BEE; text-fig. 1), the most severe extinction of deep-sea ben-
thic foraminifera of the Cenozoic (e.g., Thomas 2007). Up sec-
tion, no specimens of benthic foraminifera >500 μm have been
observed in the overlying six meters of Ypresian sedimentary
rocks and not one specimen of larger lituolids or CET taxa has
been recorded.
According to Giusberti et al. (2016), the entire benthic
foraminiferal assemblage composition (>63 μm) of the
Thanetian portion of the Forada section is dominated by
bolivinids that exploited refractory, laterally advected organic
matter. Such infaunal-dominated fauna of the Thanetian Scaglia
Rossa shows a high diversity, suggesting that seasonal to peri-
odical increases in primary productivity may have occurred in
the Belluno Basin area. The marked increase in abundance of
larger lituolids at about -40 cm below the Paleocene/Eocene
boundary at Forada (text-fig. 1) coincides with the onset of the
peak in the planktonic foraminifera Acarinina observed by
Luciani et al. (2007) and slightly precedes some of the faunal
changes detected by Giusberti et al. (2016), indicating warming
of water column and increased surface nutrient availability and
deep-water food availability.
Test morphology
The studied Paleocene lituolids show a high intraspecific mor-
phological variability (Plate 1A and figure A, available online
as supplementary material). The agglutinated tests vary from
small sphaerical to ovoidal (ranging in size from about 0.76 to
1.25 mm), and are characterized by a strongly involute coiling
of the chambers in juvenile individuals (Plate 1A, figures 1–3,
9), to larger (also exceeding 3 mm length and 1 mm wide),
mostly planispiral shells that become uncoiled in the last growth
stages, with a more or less developed uniserial chamber ar-
rangement in adult individuals (4–8). Chambers gradually in-
crease in size during the spiral stage, then they gradually reduce
the size in the uniserial stage (6), moreover, chambers are sepa-
rated by slightly arched septa, that are more visible and de-
pressed on the test surface of adult specimens (4a, 5 and 6). In
both juvenile and adult specimens, the agglutinated tests are
covered by a rough, coarse-grained surface (1–9).
Even the primary aperture morphologically changes during test
growth; it modifies from a simple straight slit-like opening in
juvenile specimens (2b) to a slightly arched slit coupled with
one or two pores in quasi-adult specimens (3b,c), finally reach-
ing a more complex morphology characterized by a higher num-
bers of elongated or circular pores (cribrate aperture), with or
without a small slit at the base in adult specimens (4c, 7).
Paleocene and Maastrichtian specimens show an apparently
identical external morphology: Cretaceous specimens have a
larger test (1.1–2.2 mm wide and 2.6–3 mm long), with a rough
surface, and chambers spirally arranged in the first growth
stages, then becoming uncoiled in the last growth stage (Plate
1B, figures 2,3) but the uncoiled portion is usually more com-
mon and well-developed in Thanetian specimens. In adult spec-
imens, the aperture is complex and formed by numerous (up to
seven), rounded or elliptical openings (Plate 1B, figure 1b).
Sectioned specimens and microstructure of the agglutinated
wall
In Paleocene and Cretaceous sectioned specimens, (Plates 2-3
and supplementary material, figures B-E), chambers are
streptospirally arranged in the initial coiling of the test, then
they become detached from the close spire according to a recti-
linear arrangement during the last growth stage, more evident in
Thanetian specimens. In the sectioned adult specimens the evo-
lution of the aperture is well manifest; it changes from a single
opening connecting the lumina in the earlier chambers to a dou-
ble or multiple opening in the last chamber (Plate 2, figures 3b,
4b, 5b; Plate 3, figures 2b-c, 3b-c).
The wall of Thanetian specimens (both juvenile and adult, Plate
2) is thick (up to about 140 mm) and complex with a typical lay-
ered microstructure formed by two distinct overlain layers (2c,
3c, 4c and 5c): an internal, thicker, carbonatic portion and an ex-
426
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Micropaleontology, vol. 64, nos. 5-6, 2018 427
Luca Giusberti, Michael A. Kaminski and Nicoletta Mancin Plate 4
PLATE 4
Elemental characterization of the agglutinated grains in a Thanetian juvenile specimen (sample BRI-37; MGP-PD 32075)
from the Forada section. Note that the external coarse-grained particles are mostly of quartz and silicates whereas the internal
portion of the wall is of calcite. Images are in back-scattered electrons (BSE).
1Longitudinal section (the same specimen of Plate 2,
image 1b) showing the elemental characterization of
the agglutinated grains. The colored crosses indicate
the spots for standardless microanalyses (A-D); on the
right the corresponding EDS spectra are reported. In
the violet spectrum (A) the very high counts of Si, to-
gether with O, indicates that the analysed grain is
quartz; similarly in the blue spectrum (B) the high
counts of Ca indicates that the grain is calcite. In the
yellow spectrum (C), the presence of Si, Al, K and O
reveals the silicatic nature (K-feldspar) of the ana-
lysed grain; finally in the green spectrum (D) the
co-occurrence of Ca, Mg, O and C is indicative of a
dolomitic grain.
2-3 Elemental maps reporting the areal distribution of Si
(image 2) and Ca (image 3) in the whole longitudinal
section of the specimen imaged above (1). Silicon and
Ca maps (here considered as discriminating elements
between quartz and calcite) show that Si is mostly
concentrated in the grains forming the external por-
tion of the agglutinated wall (the external pavement),
on the other hand, Ca homogeneously and abundantly
occurred in the internal part of the agglutinated wall.

ternal, thinner, coarse-grained pavement of mineral granules; in
the internal carbonatic portion, some particles are remains of
planktonic foraminiferal tests (3c white arrow). The aggluti-
nated grains from both layers are embedded in a homogeneous
calcareous matrix (2c; 3c, 4c and 5c). Note that these features
persist unchanged in both juvenile and adult specimens (1b, 2b;
3b, 4b, 5b) and during the test growth.
Noteworthy, in the Danian sectioned specimens (Plate 3, fig-
ures 1a-c, 2a-c) the agglutinated wall is clearly different: it is
characterized by a thick (up to about 150 mm), homogeneous,
single portion, mostly made of carbonate particles embedded
together by an abundant calcitic cement (1b-c, 2b); the largest
particles are tests of planktonic foraminifera (arrows in 1c). The
external portion of the wall, made of coarse-grained mineral
particles, a feature that is clearly visible in Thanetian speci-
mens, is here missing. It is noteworthy that these characteristics
persists unchanged during the test growth and are the same in
juvenile and adult specimens (1b-e and 2 b-d).
Likely, in the Cretaceous sectioned specimen imaged in Plate 3,
figures 3a-c, the agglutinated wall consists of a single carbonate
layer, without the external coarse grained-portion that appears
as a peculiar feature of the Thanetian specimens.
Chemical-mineralogical composition of the agglutinated
grains
In both juvenile and adult Thanetian specimens (Plates 4–6 and
supplementary material, Figures B-E), the agglutinated grains
are always selected within the test wall, in terms of chemi-
cal-mineralogical composition, size and disposition. The largest
grains are mainly of quartz (more rarely of other silicates, such
as plagioclase and k-feldspar) and are arranged toward the out-
side of the agglutinated wall to form a distinct coarse-grained
layer (a sort of external pavement) that externally covers both
the wall and the septa (Plates 4-6, Si-maps). This external pave-
ment is usually thinner with respect to the entire thickness of
the agglutinated wall (i.e., Plate 5, figure 1b; Plate 6, figure 1b)
and is formed by granules which are also selected on the basis
of their size and disposition, with the largest grains arranged
close to the outside and the smallest grains towards the inside
(Plates 4-6, Si-maps). Most of the remaining agglutinated wall
is formed by fine-grained carbonate particles (Plates 4-6,
Ca-maps), mostly of calcite and, more rarely, of dolomite or of
remains of planktonic foraminiferal tests (i.e., Plate 6, figures
1c, 2); these carbonatic grains can be less evident within the cal-
cite matrix mainly when the diagenetic process has been partic-
ularly invasive (Plate 4, figure 1; Plate 5, figure 1b).
It is interesting to note that in all the studied Thanetian speci-
mens, the agglutinated grains are strongly selected maintaining
the same mineralogical composition even in specimens col-
lected from different stratigraphic sections (e.g., uppermost
Thanetian of the Monte San Lorenzo Quarry, Friuli-Venezia
Giulia region; supplementary material, Fig. B, specimen 4).
The ability for selecting grains from the substratum to form
their test is also confirmed by the mineralogical composition of
the sediments containing the studied Thanetian lituolids (sup-
plementary material, Table A). In the Thanetian record of the
Forada section, quartz is not a dominant component (it occurs in
the sediment with percentages varying from 8.3 to 17%;
Giusberti et al. 2007); in spite of this, quartz is always selected
by the Thanetian lituolids to construct the external portion of
the test; on the contrary, phyllosilicates, that are always more
abundant in the sediments (with percentages varying from 29%
to 57%; Giusberti et al. 2007), are never observed in the studied
agglutinated tests.
Comparison with other lituolids described in the literature
Significant differences emerged when we compared the sec-
tioned Thanetian specimens from the Forada section with Upper
Cretaceous specimens attributed to Navarella Ciry and Rat
1951 and similar lituolids from other European and Italian lo-
calities and previously described in the literature (e.g., Maync
1952; 1954; Cati 1960; Sampò 1972; Riegraf 1998) (Table 1).
The investigated specimens were also directly compared with
the type material (both isolated and sectioned specimens) of the
larger lituolid “Recurvoides”manfredii Cati (1960) from the
Campanian Scaglia Rossa of Berici Hills, presently housed at
the Museo Capellini of Bologna University (Italy). We tried to
locate the original material studied by Sampò (1972) from the
Maastrichtian Scaglia Rossa of the Lessini Mountains (Veneto
region) in the collections of the Department of Earth Sciences of
Turin University, but unfortunately it seems to be lost.
In term of external morphology, Thanetian lituolid specimens
more closely resemble the genus Navarella Ciry and Rat 1951
for possessing a more developed uncoiled portion of the test in
comparison with the Campanian lituolids from Scaglia Rossa
described by Sampò (1972) and Maastrichtian and Danian spec-
imens from Forada section analyzed in this study (Table 1).
The agglutinated wall of our Thanetian specimens is typically
bi-layered, with the grains selected on the basis of their size,
shape and chemical-mineralogical composition. The larger
grains are of quartz and are arranged towards the outside of the
test wall to form a sort of rough external pavement; on the con-
trary towards the inside, the wall is formed by a more regular,
thicker calcareous portion, made of smaller carbonate grains of
calcite and more rarely of dolomite and planktonic foram-
iniferal test remains. All the agglutinated grains forming both
the external and the internal layers are cemented by calcite.
None of the investigated Thanetian specimens show an aggluti-
nated wall formed exclusively by a single layer of fine sugary
quartz grains, as reported for Navarella joaquini by Maync
(1954), nor a test wall formed by agglutinated grains of differ-
ent sizes, randomly arranged and exclusively composed by
foraminiferal tests and carbonate particles as reported by Sampò
(1972) for Maastrichtian specimens ascribed to Navarella
joaquini from the Scaglia Rossa of Verona province.
Maastrichtian and Danian specimens of lituolids recovered at
Forada are morphologically quite similar to the specimens de-
scribed by Cati (1960) as “Recurvoides”manfredii, even if the
composition of the agglutinated grains is more characteristic of
the lituolids described by Sampò (1972) as “Lituola grandis”
and “Navarella joaquini” (Table 1).
Summarizing, we conclude that the studied Thanetian lituolid
specimens cannot be attributed to the genus Navarella Ciry and
Rat 1951: they clearly differ with respect to the emended de-
scription of Maync (1954), whose specimens from Switzerland
were compared by Riegraf (1998) to Lituola westfalica
Bartenstein 1952. Riegraf (1998) also considered Navarella
joaquini and Lituola westfalica as convergent but not synony-
mous taxa, whose relationships should be explored through
careful study.
428
Luca Giusberti et al.: The bathyal larger lituolid Neonavarella n. gen. from the Thanetian Scaglia Rossa Formation, northeastern Italy

Micropaleontology, vol. 64, nos. 5-6, 2018 429
Luca Giusberti, Michael A. Kaminski and Nicoletta Mancin Plate 5
PLATE 5
Elemental characterization of the agglutinated grains in a Thanetian quasi-adult specimen (sample BRI-9; MGP-PD 32072).
Note that the compositional features of the agglutinated grains are the same previously describe for the juvenile specimen.
Images are in back-scattered electrons (BSE).
1a Longitudinal section (the same specimen of Plate 2,
image 2b) showing the elemental characterization of
the agglutinated grains.
1b Detail of the agglutinated test wall. The colored
crosses indicate the spots for standardless micro-
analyses (A-B); on the right the corresponding EDS
spectra are reported: A) quartz (in yellow) and B) al-
bite (in red). Note that silicates (e.g. K-feldspar and
plagioclase) are characterized by very high counts of
Si, together with similar counts of Al and O; the pres-
ence of Na and/or Ca, with significant counts, discrim-
inates plagioclases (albite vs. anortite), while the K
peak may be considered as indicative of K-feld-
spars.
2-4 Elemental maps reporting the areal distribution of Si
(image 2) Al (image 3) and Na (image 4) in the longi-
tudinal section of the quasi-adult specimen imaged
above (1). Silicon (here considered as indicative of
quartz and silicates) is mostly concentrated in the
grains forming the external portion of the agglutinated
wall (the external pavement), on the other hand, most
of the internal agglutinated wall is made of calcite (in
black in image 2).

Based on all the above mentioned considerations, we ascribe
the Thanetian specimens from the Forada section to a new ge-
nus of navarellid lituolid, Neonavarella n. gen. described be-
low, whereas the Maastrichtian–Danian specimens from the
Foradasectionareassignedto“Navarella” sp. (Plate 1A,
figures 8, 9; Plate 1B and Plate 3), pending further revision of
all larger lituolids from Italian Scaglia-type rocks.
SYSTEMATIC PALEONTOLOGY
We formally describe a new genus and species belonging to the
subfamily Ammobaculininae, and compare it with the
stratigraphically older and possibly ancestral genus Navarella
Ciry and Rat 1951. The following description is based on the
traditional morphology-based classification system of Loeblich
and Tappan (1987; see Loeblich and Tappan 1989) partly inte-
grated and emended by Kaminski (2014), which uses test wall
microstructure and grain composition.
Class FORAMINIFERA d’Orbigny 1826
Order LITUOLIDA Lankester 1885
Suborder LITUOLINA Lankester 1885
Family AMMOBACULINIDAE Saidova 1981
Subfamily AMMOBACULININAE Saidova 1981
Genus Neonavarella Giusberti, Kaminski and Mancin n. gen.
Type species: Neonavarella sudalpina n. gen., n. sp.
Diagnosis: Test large, initially streptospirally enrolled, invo-
lute, later uncoiling, with numerous, broad and low chambers
overlapping the earlier ones, septa strongly arched. Aggluti-
nated wall consisting of two layers: an outer layer of larger ag-
glutinated quartz grains that forms an outer pavement, and a
thicker inner calcareous layer consisting of smaller calcareous
particles in a calcareous ground mass. Cement of calcite. Aper-
ture a basal or areal slit in the early coiled stage, areal in the
later coiled stage, consisting of small circular openings in addi-
tion to the larger interiomarginal one; the uncoiled stage has a
terminal cribrate aperture.
Etymology: From the Ancient Greek, “neoz”(=new)and
“Navarella”, the Upper Cretaceous genus defined by Ciry and
Rat (1951) that is morphologically similar.
Composition of the genus: At the moment the genus is repre-
sented only by the type species Neonavarella sudalpina n. sp.
Type horizon: Thanetian (upper Paleocene) reddish and green-
ish marls and calcareous marls; calcareous nannofossil zones
CNP8-CNP11 of the scheme of Agnini et al. (2014).
Type locality: Forada Creek, Belluno province of northeastern
Italy (text-fig. 1).
Stratigraphic and geographic range: Thanetian (upper Paleo-
cene) of the Veneto region and Thanetian of San Lorenzo
Quarry, Maniago (Pordenone province, Friuli region).
Remarks: Neonavarella n. gen. differs from Navarella Ciry and
Rat 1951 in its two-layered clearly differentiated wall and youn-
ger stratigraphic range (Table 1).
Neonavarella sudalpina Giusberti, Kaminski and Mancin n. sp.
Plate 1A, figures 1–7; Plate 2; Plates 4–7; supplementary mate-
rial, figures A (pages 1–4), figures B–E, F (page 1)
“Large lituolids” of Giusberti et al. 2016, p. 219, pl. 4, figs. 14,
20.
Diagnosis: As Neonavarella is monotypic, the diagnosis of its
type species is the same as that of the genus.
Type horizon: Thanetian (upper Paleocene) reddish and green-
ish marls and calcareous marls; calcareous nannofossil zones
CNP8-CNP11 of the scheme of Agnini et al. (2014).
Type locality: Forada Creek section, Belluno province of north-
eastern Italy.
Etymology: After “Sudalpino” (Southern Alps), the name of the
sector of the Alpine Chain in which the new taxon was discov-
ered.
430
Luca Giusberti et al.: The bathyal larger lituolid Neonavarella n. gen. from the Thanetian Scaglia Rossa Formation, northeastern Italy
PLATE 6
Elemental characterization of the agglutinated grains in a Thanetian adult specimen (thin section MGP-PD 32072, sample BRI-9).
Note that also in the adult specimen the compositional features of the agglutinated wall do not change.
Images are in back-scattered electrons (BSE).
1a Longitudinal section showing the elemental charac-
terization of the agglutinated grains.
1b-c Details of the agglutinated wall. The colored crosses
indicate the spots for standardless microanalyses
(A-C); on the right the corresponding EDS spectra are
reported: A) calcite (in yellow); B) quartz (in blue)
and C) barite (in red).
2-4 Elemental maps reporting the areal distribution of Ca
(image 2), Si (image 3) and Al (image 4) in the longi-
tudinal section of the adult specimen imaged above
(1). Silicon (here considered as indicative of quartz
and silicates) is mostly concentrated in the grains
forming the external portion of the agglutinated wall
(the external pavement), on the other hand, most of the
internal agglutinated wall is made of calcite (image 2).
Note the remains of planktonic foraminifera (arrows)
within the carbonate portion of the agglutinated wall.

Micropaleontology, vol. 64, nos. 5-6, 2018 431
Luca Giusberti, Michael A. Kaminski and Nicoletta Mancin Plate 6

Repository: The type series (MGP-PD 32071-32073, 32075-
32088) is housed in the micropaleontological collections of the
Museum of Geology and Paleontology of the University of
Padova (Italy). Additional topotypic specimens are housed in
the personal collection of one of the authors (L.G.) at the De-
partment of Geosciences of Padova University (Italy), and in
the collections of the European Micropalaeontological Refer-
ence Centre, at Micropress Europe, Kraków Poland; such speci-
mens are labelled with the sample number assigned during the
sampling of the section.
Designated holotype: The isolated specimen illustrated in Plate
7 and registered as MGP-PD 32068.
Paratypes: Figured paratypes (MGP-PD 32071-32073, 32075-
32081) are in Plate 1A, figures 1–7; Plate 2; Plates 4–6; and
supplementary material, figs. A (pages 1–4) and B–F (page 1).
Unfigured paratypes are registered as MGP-PD 32082-32087.
Description: High intraspecific morphological variability from
juvenile to adult specimens. Juvenile specimens generally
smaller in size (ranging from ~0.76 to 1.25 mm), ovoidal to
subspherical with chambers strictly coiled. Early chambers
streptospirally, then planispirally arranged (7–8 chambers in the
last whorl) passing to uncoiled in the adult larger specimens
(also exceeding 3 mm length and 1 mm wide). Uncoiled portion
of the test usually less well-developed, consisting of 3–4 small
chambers. Primary aperture that morphologically changes dur-
ing test growth: slit from almost rectilinear in the juvenile speci-
mens to markedly arched. In adult individuals, the apertural slit
is associated with small circular pores (up to a maximum of 4
pores); a truly cribrate aperture (with 6–7 small rounded pores)
is less frequent in the adult uncoiled specimens. Agglutinated
wall typically bi-layered, formed by an external coarse-grained
layer of silicilastic grains mainly of quartz and an inner thicker
layer of calcareous grains (also biogenic remains, as foram-
iniferal tests). Grains held together by calcitic cement.
432
Luca Giusberti et al.: The bathyal larger lituolid Neonavarella n. gen. from the Thanetian Scaglia Rossa Formation, northeastern Italy
PLATE 7
Neonavarella sudalpina n. gen., n. sp. Holotype. Slide MGP-PD 32068. Sample BRI-466. Scaglia Rossa Formation, Thanetian,
Forada section, northeastern Italy. a) lateral view, b) apertural view showing a slit-like aperture coupled with three pores

Remarks: quite similar to Maastrichtian–Danian specimens
from the Forada section (“Navarella” sp.) from which
Neonavarella sudalpina n. sp. differs in the bi-layered wall
structure (Table 1).
Stratigraphic and geographic range: Thanetian (upper Paleo-
cene; calcareous nannofossil zones CNP8-CNP11 of Agnini et
al. 2014) of the Veneto region (Italy) and uppermost Thanetian
(calcareous nannofossil zone CNP11 of Agnini et al. 2014) of
San Lorenzo Quarry, Maniago (Pordenone province, Friuli re-
gion, Italy). The taxon becomes extinct at the Paleocene/
Eocene boundary, in coincidence with the BEE (benthic
foraminiferal extinction event).
Paleoenvironment: Basinal setting at middle-lower bathyal
depth (~1000–1500 m). The new taxon is a minor component of
the >63 μm fraction benthic foraminiferal assemblage, strongly
dominated by bolivinids. Neonavarella sudalpina n. gen., n. sp.
is associated in the >500 μm fraction with deep-water taxa such
as clavulinids, Cicicidoides, anomalinids and trochamminids.
This assemblage includes also some characteristic Paleocene
extinction taxa, such as Clavulinoides globulifera,Cibicidoides
dayi and C. velascoensis (CET, Cosmopolitan Extinction Taxa
of Giusberti et al. 2016).
CONCLUSIONS
A direct comparison among Maastrichtian, Danian and
Thanetian specimens of larger lituolids from the Forada section
(Belluno basin, northeastern Italy) permitted the assignment of
the Thanetian lituolids to a new genus here formally designated
as Neonavarella n. gen., a singleton taxon (sensu Foote 2000)
morphologically similar to Navarella Ciry and Rat 1951, but
characterized by a peculiar bi-layered agglutinated wall.
The older Maastrichtian-Danian specimens from the Forada
section are here ascribed to “Navarella” sp.; nonetheless the re-
lationships between Navarella sensu Ciry and Rat 1951 and the
specimens of “Navarella” from the Upper Cretaceous Scag-
lia-type rocks of northeastern Italy need to be more fully ex-
plored.
This study also allows the stratigraphic range of larger bathyal
navarellid lituolids, previously thought to be limited to the Up-
per Cretaceous, to be extended into the Paleocene. Finally, we
demonstrate the importance of compositional and micro-
structural studies based on sectioned specimens for establishing
the taxonomic significance of the wall structure in agglutinated
foraminifera.
ACKNOWLEDGMENTS
The authors deeply thank Roberto Barbieri and Carlo Sarti for
access to type series of Recurvoides manfredii Cati housed at
the Museum Capellini of Bologna University. We also ac-
knowledge the Arvedi’s Laboratory (CISRiC, University of
Pavia) for SEM-EDS analyses and Stefano Castelli (Diparti-
mento di Geoscienze, University of Padova) for accurate photo-
graphic documentation of some specimens illustrated in this
paper. L.G. was financially supported by the University of
Padova ex 60% Fornaciari E.; MAK acknowledges support
from a start-up grant from the College of Petroleum and
Geosciences, KFUPM. N.M. was financially supported by
“Fondo Ricerca Giovani” FRG-2016 and FFARB funds (resp.
N.M., University of Pavia).
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Supplementary materials (Available on-line)
FIGURE A
Intraspecific morphological variability of the studied lituolids
picked from the whole upper Maastrichtian, Danian and
Thanetian stratigraphic record of the Forada section. Collected
specimens vary from small juvenile individuals with a simple
slit-like aperture and a strongly involute coiling of the test to
larger adult individuals with a complex aperture and a test that
becomes uncoiled in the last growth stages. Pages 1-4:
Thanetian specimens (Neonavarella n. gen.). Page 5: Danian
specimens (“Navarella” sp.). Page 6: Maastrichtian specimens
(“Navarella” sp.).
Page 1: all specimens from slide MGP-PD 32078 (except the
specimen n. 9 from section MGP-PD 32075. Page 2: all speci-
mens from slide MGP-PD 32079 (except the specimens n. 5 and
9 from MGP-PD 32075 and the specimen 7 from MGP-PD
32071). Page 3: all specimens from slide MGP-PD 32080 (ex-
cept the specimen 8 from MGP-PD 32071 and the specimens n.
10 and 12 from MGP-PD 32073). Page 4: all specimens from
slide MGP-PD 32080 (except the specimen 16 from MGP-PD
32071 and the specimens n. 17 and 18 from MGP-PD 32073).
Page 5: specimens 1, 3 and 5 from MGP-PD 32074, specimens
4 from slide with sample number RF +670C. Page 6: specimens
1, 2, 3, 7 are housed in slides labeled “RF-24", specimen 4, 5
from slide MGP-PD 32089, specimens 6 and 9 sectioned
(MGP-PD 32069) and specimen 8 sectioned (MGP-PD 32070).
FIGURES B-E
Sectioned specimens of Thanetian lituolids (Neonavarella
sudalpina n. gen., n. sp.) from the Forada and Maniago sections,
showing the microstructure of the agglutinated wall formed by
two distinct layers and the chemical-mineralogical composition
of the agglutinated grains of both adult and juvenile specimens.
Note that the microstructural and compositional features of the
agglutinated tests persist unchanged in both juvenile and adult
specimens and during the test growth. Similarly, no evident dif-
ferences merged in individuals picked from different strati-
graphic layers.
Figure B: thin section MGP-PD 32072 (including the specimen
4, coming from the uppermost Thanetian of Monte San Lorenzo
Quarry section, Friuli region), Figure C: thin section MGP-PD
32071, Figure D: thin section MGP-PD 32075, Figure E: thin
section MGP-PD 32073. Thin section MGP-PD 32071 in Fig-
ure C includes also four sectioned specimens of Pliocene
Colominella published by Mancin and Kaminski 2017 - Sys-
tematic updates of the agglutinated foraminiferal genus
Colominella Popescu 1998: insights from sectioned specimens.
Geologica Carpathica 68(2), 109-118.
FIGURE F
Images of Thanetian, Danian and Maastrichtian sectioned speci-
mens prepared as thin sections and analyzed with a light polar-
ized microscope. Page 1: specimens from section MGP-PD
32073. Page 2: specimen from section MGP-PD 32069. Page 3:
specimen from section MGP-PD 32074.
TABLE A
Mineralogical characterization (RDX data) of the sediments
containing the studied lituolids (from Giusberti et al. 2007).
434
Luca Giusberti et al.: The bathyal larger lituolid Neonavarella n. gen. from the Thanetian Scaglia Rossa Formation, northeastern Italy