Cretaceous Research 24 (2003) 1–22

Calcareous nannofossils from the Aptian–Lower Albian of

southeast France: palaeoecological and biostratigraphic implications
Jens O. Herrle a*, Jörg Mutterlose b
a
Institut für Geowissenschaften, Universität Tübingen, Sigwartstrasse 10, D-72076 Tübingen, Germany
b
Institut für Geologie, Mineralogie und Geophysik, Ruhr-Universität Bochum, Universitätsstrasse 150, D-44780 Bochum, Germany
Accepted 2 September 2002

Abstract

The Aptian–lower Albian succession of the Vocontian Basin (SE France) consists of marine hemipelagic sediments including
several black shale horizons. The latter are partly of regional and partly of global distribution. This sedimentary succession records
the nannoplankton evolution of the Aptian–early Albian interval and thus provides an excellent opportunity to calibrate the
calcareous nannofossil record with Tethyan ammonite and planktic foraminiferal biostratigraphy. The calcareous nannofossil
biostratigraphy presented in this paper supports previous zonations, but it also provides a much higher resolution and thus improves
the correlation of different black shale horizons on a supraregional scale. Up to 23 major (supraregionally significant) and minor
(regionally significant) first and last occurrences of calcareous nannofossil taxa are recognized. Nannoconid abundances decrease
rapidly in the upper Lower Aptian (nannoconid crisis I, NCI) and in the middle Upper Aptian (nannoconid crisis II, NCII). Both
decreases correlate with carbonate–platform drowning events. The upper Lower Aptian interval above the NCI is characterized by
high abundances of large specimens of Assipetra infracretacea and Rucinolithus terebrodentarius probably of supraregional
significance. The uppermost Aptian–Lower Albian is characterized by high abundances of the calcareous nannoplankton taxon
Repagulum parvidentatum, reflecting boreal influence on the Tethyan Realm. This suggests a temporary decrease in surface-water
temperatures in the Vocontian Basin.
 2003 Published by Elsevier Ltd.

Keywords: Calcareous nannofossils; Biostratigraphy; Early Cretaceous; Aptian; Albian; France; Vocontian Basin

1. Introduction warmer than at present (Savin, 1977; Erbacher

The mid-Cretaceous, lasting from the Aptian to the
Turonian (120–90 Ma; Gradstein et al., 1995), has been
characterized as a time of global warmth when the
Earth’s climate was in an extreme ‘greenhouse’ mode.
This is indicated by high atmospheric pCO2 (Berner
et al., 1983; Barron & Washington, 1985), elevated
average temperatures (e.g., Barron et al., 1995), a rela-
tively high-stand sea-level (Haq et al., 1987), and a
reduced latitudinal temperature gradient as compared
to the Holocene (e.g., Spicer & Cornfield, 1992; Huber
et al., 1995). Oxygen isotope data from benthic foramin-
ifera indicate that the deep ocean was significantly
* Corresponding author: J.O. Herrle, ETH Zurich, Geological
Institute, Soneggstrasse 5 NO G28, Zurich 8092, Switzerland.
Tel.: 0041-(0)1-632-3694; Fax: 0041-(0)1-632-1080
E-mail address: Jens.Herrle@erdw.ethz.ch (J.O. Herrle).
0195-6671/03/$ - see front matter  2003 Published by Elsevier Ltd.
doi:10.1016/S0195-6671(03)00023-5
et al., 2001). Among others, these observations have
been taken as evidence that deep water formation
occurred at low latitudes within the Tethyan Ocean
(Brass et al., 1982), a conclusion that has been verified
by oceanic general circulation models (Barron &
Peterson, 1990).
The specific conditions in mid-Cretaceous times
favoured the deposition and burial of organic, carbon-
rich sediments subsequently preserved as black shales
(e.g., Schlanger & Jenkyns, 1976; Arthur & Premoli
Silva, 1982). Black shales of both regional and supra-
regional significance are most common in the western
Tethys and less abundant in the Atlantic and Pacific
oceans (e.g., Erba et al., 1989; Arthur et al., 1990;
Bréhéret, 1997). Supraregionally distributed black
shales, which are encountered in all major oceans, are
commonly referred to as Oceanic Anoxic Events (OAEs;
2 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
Schlanger & Jenkyns, 1976; Arthur et al., 1990;
Bralower et al., 1993, 1994; Erbacher et al., 2001; Herrle
et al., 2003). The OAEs (e.g., the late early Aptian OAE
1a and early Albian OAE 1b) are suggested to have been
accompanied by prominent biotic changes reflected in
calcareous nannofossils, planktic and benthic foramin-
ifera, radiolarians, belemnites, and ammonites (e.g.,
Erba, 1994; Erbacher & Thurow, 1997; Cecca, 1998;
Mutterlose, 1998, Premoli Silva et al., 1999; Herrle et al.,
2003), widespread drowning of shallow carbonate plat-
forms (Weissert et al., 1998; Lehmann et al., 2000;
Strasser et al., 2001), oceanic volcanic eruptions in the
Pacific, sea-level changes, and the onset of Cretaceous
greenhouse conditions (Larson, 1991). These relation-
ships of black shales and biotic and palaeoclimatic
changes in the mid-Cretaceous require a biostrati-
graphic framework in order to allow accurate dating
and correlation of the various events recorded.
The main aim of this paper is to present a high-
resolution calcareous nannofossil biostratigraphy cali-
brated to the Tethyan ammonite and planktic
foraminiferal biostratigraphies of the Aptian–Lower
Albian succession. In addition, it aims to discuss the
distribution of the typical Tethyan and Boreal nanno-
plankton taxa during the Aptian–early Albian. A com-
posite section in the Vocontian Basin (SE France),
which is rich in calcareous nannofossils and black shale
horizons, was selected as a reference section for this
study.
2. Geologic and palaeogeographic setting of the
Vocontian Basin (SE France)
The Vocontian Basin is located between the Vercors
Massif in the north, the river Rhône in the east, the
Ventoux-Lure Axis, and the Nice-Castellane Arc in the
south (Fig. 1). It corresponds to the Diois and Baronnies
regions and the Verdon valley of the Southern Subalpine
Ranges.
The Mesozoic sediments of the Vocontian Basin
are characterized by marlstone-dominated (Upper
Bajocian–Lower Oxfordian, Middle Valanginian, Lower
Aptian–Lower Cenomanian) and limestone-dominated
(Upper Oxfordian–Lower Valanginian, Upper
Valanginian-lowermost Aptian, Cenomanian-Turonian;
Arnaud & Lemoine, 1993) intervals. In the upper Lower
Aptian, there is a sharp decrease in the carbonate con-
tent and a monotonous succession of marly hemi-
pelagic sediments. These sediments constitute the
Marnes Bleues Formation (Flandrin, 1963), which is up
to 750 m thick. This unit is marked by pale- and dark-
bedded marlstones intercalated with marly limestone,
limestone, and a large number of black shale horizons
(Bréhéret, 1988, 1997). The entire succession is charac-
terized by cyclic deposits (e.g., Bréhéret, 1994; Kößler
et al., 2001) occasionally interrupted by turbidites and
slumps (Bréhéret, 1997). Calcareous sediments become
more abundant again in the Upper Albian.
The Vocontian Basin was the hemipelagic part of the
Mesozoic Dauphinois Basin during mid-Cretaceous
time and formed a more or less intracratonic structure
(Baudrimont & Dubois, 1977). From the Late Jurassic
onwards throughout the Early Cretaceous it was situ-
ated at a palaeolatitude of 25–30( North (Savostin et al.,
1986; Hay et al., 1999). It was open towards the Tethys
to the east and surrounded by slopes and platforms with
hemipelagic facies intercalating with shallow-water car-
bonates (Fig. 2). In latest Hauterivian–early Aptian
time the Vocontian Basin was surrounded by Urgonian
carbonate (coral-rudistid) platforms, i.e., the Vercors
Platform in the north and the Provence Platform in
the south (Arnaud-Vanneau & Arnaud, 1990, 1991). To
the east, the connection of the Vocontian Basin to the
Tethys Ocean has been obliterated by the thrust fault
system of the Alpine Penninic units (Arnaud & Lemoine,
1993).
Based on floral and faunal evidence, the depth of
water in the Vocontian Basin is estimated to have been
several hundred metres (Wilpshaar & Leereveld, 1994;
Wilpshaar et al., 1997), although some authors have
suggested that the depth exceeded 2000 m (Cotillon &
Rio, 1984). However, in view of the overall composition
of the dinoflagellate cyst assemblages (Wilpshaar &
Leereveld, 1994), and according to sedimentological
criteria (De Boer, 1983), this seems unlikely.
3. Material and methods
The Aptian–Lower Albian succession (340 m) dis-
cussed is a composite of seven sections covering different
intervals of time (Fig. 3). The individual sections were
correlated using the basin-wide marker horizons
(Faisceau Nolan, Faisceau Fromaget, Délit Calcaire 4,
Niveau Kilian, Haut Noire 7 and Niveau Paquier; see
below) described by Bréhéret (1997).
Sample intervals vary from 0.8 to 1 m. A total of 383
samples well prepared for their nannofossil content
using standard smear-slide techniques (e.g., Bown &
Young, 1998). Biostratigraphically important taxa and
abundances of nannoconids and the typical boreal cal-
careous nannoplankton taxon Repagulum parvidentatum
(e.g., Mutterlose & Wise, 1990) were documented semi-
quantitatively. Three classes were used to denote abun-
dances. These are defined as follows: common, >1
specimen/field of view (FOV); frequent, 1 specimen/2–10
FOV; rare, 1 specimen/>11 FOV.
In addition, high-resolution studies at sample inter-
vals of 2–10 cm were carried out (339 samples) on
selected black shale and carbonaceous intervals (Niveau
Goguel to Niveau Blanc; Niveau Kilian, Niveau
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
Fig. 1. Map showing major geological features in southeast France (modified after Lorenz, 1980) and the location of the area studied in the
Vocontian Basin.
Paquier; see Fig. 3). The samples were prepared using
the method described by Geisen et al. (1999). At least
300 calcareous nannofossil specimens were counted in
random traverses of each slide using an Olympus BX50
photomicroscope at a magnification of 1250.
Taxa considered here are listed in the Appendix.
Bibliographic references for these taxa are given in
Perch-Nielsen (1985) and Bown et al. (1998). The slides
are housed in the Institute for Geosciences in Tübingen
(Germany). All data presented and detailed range charts
of the intervals studied at high resolution are available in
the PANGAEA database (http://www.Pangaea.de).
4. Localities and lithology
Seven sections were studied with respect to their
contents of calcareous nannofossils. These are Serre
3
Chaitieu, Gaubert, Tarendol, Pré-Guittard, Les
Oustaus, l’Arboudeysse, and Col de Palluel (Figs. 3 and
4). The local nomenclature for individual black shale
horizons (Niveau Goguel, Niveau Noirs, Niveau Noire
Calcaire 2, Niveau Fallot, Niveau Jacob, Niveau Kilian,
Haut Noirs, Niveau Paquier, and Niveau Leenhardt)
and carbonaceous intervals (Niveau Clairs, Niveau
Blanc, Faisceau Nolan, Faisceau Fromaget, Délits
Calcaires) follows Bréhéret (1983, 1991) and Bréhéret
(1997). The black shale horizons are dark and lami-
nated. The Niveau Goguel (OAE 1a) and the Niveau
Paquier (OAE 1b) have the additional features of a
paper shale (Bréhéret, 1997). The stratigraphic ranges of
the sections are given in Fig. 3. All sections studied apart
from that at l’Arboudeysse have been recently described
in detail by Bréhéret (1997). A geographic map with
locations of the sections is given in Fig. 4 and a
4 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22

Fig. 2. Paleogeographic reconstruction of the Vocontian Basin for the late Aptian–early Albian showing the sections studied. SC, Serre Chaitieu;
GA, Gaubert; TA, Tarendol; PG, Pré-Guittard; OUS, Les Oustaus; PL, Col de Palluel with l’Arboudeysse. Modified after Arnaud & Lemoine
(1993).

lithological column of the coraposite section is provided
in Fig. 5.
4.1. Serre Chaitieu section
This section is situated about 1 km south of
Lesch-en-Diois, on the southern and northern hillsides
of the Serre Chaitieu: topographic map (TM) 25
Luc-en-Diois, No. 3238 Ouest, Série Bleues, Lambert III
coordinates X: 853 213, Y: 3259 250. It encompasses
Lower–Upper Aptian deposits including the interval
below the Niveau Goguel black shale to the Faisceau
Nolan 3 (FN3). There are about 140 m of bioturbated
Aptian marlstones and intercalated black shale layers,
bioturbated marly limestones and limestones (Fig. 5).
The succession shows bedding rhythms of numerous
bioturbated pale and dark marlstone beds. The lower
part of the section is characterized by five black shale
horizons (laminated, non-bioturbated) 2.5 m thick,
known as the Niveau Goguel (Bréhéret, 1988).
Approximately 13 m above this level, a basin-wide marly
limestone marker bed is present, namely the Niveau
Blanc (Friès et al., 1985). About 15 and 35 m above the
Niveau Blanc two marly black shale horizons, the
Niveau Noir, NN1–4, and the Niveau Noir Calcaire 2,
NNC2, occur (Fig. 5). NN1 marks a basin-wide colour
change from pale to dark marlstones (Friès & Rubino,
1990). In the lower Upper Aptian, ten organic-rich
layers, known as the Niveau Fallot (FA1–4; Bréhéret,
1997) are present. The entire Niveau Fallot is 20 m thick.
The upper part of the succession is characterized by
marlstones intercalated with sandy turbidites, followed
by marly limestones and limestones known as the
Faisceau Nolan (FN1–3). The upper part of the
Faisceau Nolan (FN4-8) is missing owing to slumping
(Bréhéret, 1997).
4.2. Gaubert section
This section is situated about 1 km north of
Baudinard, on the southern hillside of l’Adret de
Baudinard: TM 25 La Motte-du-Caire, No. 3339 Est,
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
Fig. 3. Summary of the stratigraphical ranges of the sections studied in southeast France with the most important key beds indicated (after Bréhéret,
1997). N., Niveau; F., Faisceau; OAE, Oceanic Anoxic event.
Lambert III coordinates X: 904 525, Y: 3228 800. It
comprises Upper Aptian deposits from the Faisceau
Nolan (FN) to the Faisceau Fromaget (FF). About 50 m
of marlstones with marly limestones and limestone beds
have been investigated (Fig. 5). The lower 25 m (FN3–8)
and the upper 5 m (FF1–2) are characterized by an
alternation of marly limestones and limestones. The
middle part, having a thickness of 20 m, is dominated by
marlstones and marly limestones.
4.3. Tarendol section
This section is situated about 2 km southwest of
Tarendol, on the eastern hillside of St. Etienne: TM 25
Rémuzat, No. 3139 Ouest, Série Bleues, Lambert III
coordinates X: 840 200, Y: 4913 800. The exposure
encompasses the Upper Aptian from 2 m below FF3 to
the Délits Calcaire 4 (DC4). There are about 4 m of
marlstones, punctuated by marly limestones, limestones
5
and black shale layers (Fig. 5). The lower part is
represented by alternating marl and limestone beds
corresponding to the upper part of FF3–7. A slump is
intercalated between beds FF6 and FF7. Approximately
2.5 m above the Faisceau Fromaget two black shale
layers rich in ammonites (Hypacanthoplites jacobi) are
present, namely the Niveau Jacob (Bréhéret, 1983). The
middle part shows bedding rhythms of numerous pale
and dark beds, intercalated by three limestone beds
known as DC2–4. This levels have a basin-wide distri-
bution and are partly rich in aucellinids (Bréhéret, 1997).
4.4. Pré-Guittard section
This section is situated on the southeast flank of the
Serre Sablon, about 800 m southeast of Pré-Guittard:
TM 25 Dieulefit, No. 3138 Ouest, Lambert III coordi-
nates X: 836 800, Y: 3248 825. It encompasses the
Upper Aptian deposits including the interval of DC4
6 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
Fig. 4. Detailed location map of the sections studied.
to a marly limestone bed 2 m above the Niveau Kilian
(Fig. 5). About 8 m of marlstones, marly limestones,
and black shales have been investigated. The interval
from DC4 to the upper part of the black shale horizon
Niveau Kilian is characterized by alternating pale and
dark beds. The Niveau Kilian black shale is 74 cm
thick. Aucellinids are abundant in the upper part of
this black shale.
4.5. Les Oustaus section
This section is situated about 500 m northeast of the
hill Les Oustaus, about 3 km southwest of Esparron:
TM 25 Serres/Veynes, No. 3338 Ouest, Lambert III
coordinates X: 881 850, Y: 3242 775. It covers Upper
Aptian–Lower Albian deposits from 2 m above the
Niveau Kilian to the Haut Noir 7 (HN7). There are
about 39 m of marlstones, marly limestones and black
shales (Fig. 5). Several black shale horizons known as
HN1–7 (Bréhéret, 1997) are intercalated with pale and
dark marlstones. Above the HN1 a basin-wide colour
change within the marlstones from pale to dark is
present (Friès & Rubino, 1990). This is followed by
marlstones with several intercalated horizons of
phosphatic concretions (Bréhéret, 1997).
4.6. L’Arboudeysse section (Col de Palluel)
This section is situated 700 m southwest of
l’Arboudeysse, about 900 m east of the Col de Palluel:
TM 25 Rosans, No. 3239 Ouest, Serie Bleues, Lambert
III coordinates X: 854 850, Y: 3238 825. It encompasses
the interval from the HN7 up to 4 m above the Niveau
Paquier of the Lower Albian. About 14 m of marlstones
are exposed (Fig. 5). They are punctuated by up to
30-cm-thick black-shale layers, HN7–8. The Niveau
Paquier black shale is 1.63 m thick and rich in ammo-
nites, with abundant leymeriellids. In the upper part of
this shale a carbonate layer rich in nannoconids up to
3 cm thick is present (Bréhéret et al., 1986; Herrle, 2002).
This so-called ‘
-bed’ (Bréhéret, 1983) is a distinctive
marker horizon throughout the Vocontian Basin.
4.7. Col de Palluel section
This section is situated 700 m northeast of the Col de
Palluel: TM 25 Rosans, No. 3239 Ouest, Serie Bleues,
Lambert III coordinates X: 854 225, Y: 3239 300. It
covers the Lower Albian interval from 4 m above the
Niveau Paquier through the Niveau Leenhardt to the
Haut Noir 21 black shales (Fig. 5). There are about 43 m
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
of marlstones and black shales. The succession shows
discrete pale and dark bedding. In the lower part black
shale beds HN13–16 are present (Bréhéret, 1994). The
middle part is characterized by a basin-wide black shale
horizon, the Niveau Leenhardt (Bréhéret et al., 1986). In
the upper part of the section up to six black shale
horizons (HN18–21) occur.
5. Results
5.1. Calcareous nannofossil biostratigraphy of the
Vocontian Basin
A comprehensive nannoplankton zonation scheme
for Lower Cretaceous low-latitude sequences, including
the Aptian–Albian succession in the Vocontian Basin,
was proposed by Thierstein (1971, 1973). However, in
recent years the calcareous nannofossil biostratigraphy
for this interval has been continuously modified and
improved by calibration with planktic foraminiferal and
ammonite biostratigraphies and magnetostratigraphy
(Erba & Quadrio, 1987; Erba, 1988; Bralower et al.,
1993, 1994, 1995; Bischoff & Mutterlose, 1998; Bown
et al., 1998; Erba et al., 1999; Jeremiah, 2001).
Calcareous nannofossils are common to abundant
and fairly well to well preserved in the sections studied
(see also Thierstein, 1973). Twelve major (supraregion-
ally significant) and 11 minor (regionally significant)
first and last occurrences (FOs and LOs) have been
recognized (Fig. 5). By calibrating nannofossil FOs
and LOs to regional and supraregional black shale
marker beds, the NC zonation (Roth, 1978; Bralower
et al., 1993, 1995) is correlated with the lithostratigra-
phy. This zonation scheme has been selected because
of its applicability to pelagic deposits and con-
tinental margins in all ocean basins (Bralower et al.,
1993).
5.1.1. Chiastozygus litterarius Zone (NC6), early
Aptian
This zone is defined as the interval from the FO of
Hayesites irregularis to the FO of Eprolithus floralis. The
lower part of the section corresponds to the upper part
of the Chiastozygus litterarius Zone NC6, which includes
the Niveau Goguel black shales (OAE 1a) up to 1 m
(FO of E. floralis, sample NG1) above this horizon
(Fig. 5).
5.1.2. Rhagodiscus angustus Zone (NC7), late Aptian
Zone NC7 is defined as the interval from the FO
of Eprolithus floralis to the FO of Prediscosphaera
columnata (Fig. 5).
5.1.3. Subzone NC7A
This subzone is defined as the interval from the FO of
E. floralis to the LO of Micrantholithus spp. The succes-
7
sion from 1 m above the Niveau Goguel black shales
(FO of E. floralis, sample NG1) to 7.7 m below the
Niveau Noire Calcaire 2 (NNC2; LO Micrantholithus
obtusus, sample PK133) in the Serre Chaitieu section
corresponds to this subzone (Fig. 5). Several FOs were
observed. From bottom to top these are the FOs of
Radiolithus planus (3.7 m above the Niveau Goguel,
sample SSC24), Eprolithus apertior (15 m above the
Niveau Goguel, sample SSC31), Braarudosphaera afri-
cana and Braarudosphaera hockwoldensis (within the
Niveau Blanc, samples PSC7, PSC12), and Eprolithus
varolii and Corollithion acutum (both 3 m above the
Niveau Blanc, samples PSC31, PSC32).
5.1.4. Subzone NC7B/NC7C
Subzones NC7B/NC7C are represented by the inter-
val from the LO of Micrantholithus spp. to the FO of
Predicosphaera columnata. The stratigraphic interval
from 7.7 m below the NNC2 (LO Micrantholithus
obtusus) to 3 m above the Délits Calcaire 2 (DC2)
corresponds to these subzones (Fig. 5). The FO of
Rhagodiscus achylostaurion, which characterizes the
base of NC7C, is not observed in the Serre Chaitieu
section. Therefore, both subzones were combined in this
study.
In the middle part of these subzones the following
species first appear: Lapideacassis mariae (5.5 m below
NNC2, sample PK113), Prediscosphaera sp. (within
NNC2, sample PK76), Orastrum perspicuum (14 m
above Niveau Fallot 4, sample SC East 209), Lapi-
deacassis glans (17 m above Niveau Fallot 4, sample SC
East 199), and Prediscosphaera spinosa (sample LF112/
3/98, within Faisceau Fromaget 5 (FF5), 20 m below the
FO of P. columnata).
5.1.5. Prediscosphaera columnata Zone (NC8), late
Aptian–early Albian
Zone NC8 represents the interval from the FO Pre-
discosphaera columnata to the FO of Axopodorhabdus
albianus. The P. columnata Zone begins 3 m above DC2
and comprises the upper part of the sections studied
(upper part of the Tarendol, Pré-Guittard, Les Oustaus
and Col de Paluell sections). Axopodorhabdus albianus
does not occur in the interval investigated (Fig. 5).
5.1.6. Subzone NC8A
The subzone NC8A is defined as the interval from the
FO of Prediscosphaera columnata to the FO of Hayesites
albiensis. It comprises the stratigraphical interval 3 m
above DC2 (FO of P. columnata, sample LF151/18/98)
to 3 m above the Niveau Kilian black shale (Fig. 5).
Several FOs were found in this subzone, including the
FOs of Eiffellithus hancockii 4.6 m above DC2 (sample
LF161/2/98), Cribrosphaerella ehrenbergii 1.5 m below
Délits Calcaire 3 (DC3; sample LF173/18/98), Heli-
colithus trabeculatus 1.9 m (sample PG218/2/98) and
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 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22 9

Seribiscutum primitivum 1 m below the Niveau Kilian
black shale (sample PG224/3/98), and Arkhangelskiella
stenostaurion within it (sample PG226/13/98). The FO of
Hayesites albiensis occurs 3 m above it (sample OUS15).
5.1.7. Subzone NC8B
Subzone NC8B is defined as the interval from the FO
of Hayesites albiensis to the FO of Tranolithus orionatus.
It comprises the interval from 3 m above the Niveau
Kilian black shale (FO of H. albiensis) to the top of the
succession investigated (Fig. 5). Within it, Sollasites
falklandensis occurs 6.9 m below the Niveau Paquier
(sample MP2–8). The occurrence of this species is fol-
lowed by the FOs of Kokia sp. and Placozygus cf.
P. fibuliformis (samples MP2-21, MP2-23), the latter
occurring only rarely.
5.2. Distribution of selected calcareous nannofossil taxa
The calcareous nannofossil assemblages recovered
are of high diversity (mean, 31 taxa per sample). Some
taxa (Biscutum constans, Discorhabdus ignotus,
Lithraphidites carniolensis, Rhagodiscus asper, Watznau-
eria barnesae and Zeugrhabdotus erectus) are abundant
to common, varying from 4 to 41% (mean values)
throughout the sections (Herrle, 2002). Nannoconus spp.
and Repagulum parvidentatum, however, show charac-
teristic fluctuations. Nannoconus spp. are common
within the Niveau Blanc, Niveau Noir 4, Niveau Noire
Calcaire 2, the interval from 6 m above Niveau Noire 4
to the Faisceau Nolan 2, and within the Niveau Paquier
black shale (Fig. 5).
Repagulum parvidentatum is rare to frequently abun-
dant in an interval above the Niveau Fallot 4 to Délits
Calcaire 4 (Fig. 5). A significant increase in numbers of
this species occurs in the uppermost Aptian, beginning
below the Niveau Kilian black shale (11.2%; mean;
2.8%) extending into the Lower Albian Niveau Paquier
black shale where abundances of 18.7% were recorded
(Herrle, 2002).
The lower part of the Serre Chaitieu section, which
covers the interval from the Niveau Goguel to
the Niveau Blanc, is characterized by the absence of
Nannoconus spp. and higher percentages of large A.
infracretacea/R. terebrodentarius (12%; Fig. 9). Assipetra
infracretacea reaches up to 7% and R. terebrodentarius
up to 5% in this interval. Above the Niveau Blanc
abundances of <1% were recorded (Fig. 9). A rapid
increase of Nannoconus spp. was observed within the
Niveau Blanc.
6. Discussion
6.1. Biostratigraphy
The Aptian–early Albian succession of the Vocontian
Basin provides a complete record of the calcareous
nannofossil evolution and, hence, providing an excellent
opportunity to calibrate the nannofossil signal to the
Tethyan ammonite and planktic foraminiferal biostrati-
graphies. In the last three decades the Lower Cretaceous
of southeast France has been the subject of numerous
studies aimed at developing biostratigraphic zonations
using calcareous nannofossils (e.g., Thierstein, 1971,
1973; Barrier, 1977a,b; Deres & Achéritéguy, 1980;
Bralower et al., 1993; Bergen, 1994; Kennedy et al.,
2000). The calcareous nannofossil zonation presented in
this paper improves previous zonation schemes by pro-
viding a much higher resolution. Twenty-three first and
last occurrences of calcareous nannofossils can be cor-
related with the ammonite and planktic foraminiferal
zonation scheme (Fig. 5). Twelve first occurrences are of
supraregional significance, whereas another 11 are prob-
ably of regional significance. Important taxa in the
sections studied are shown in Figs. 6–8.
The lower Aptian part of the succession. The order of
succession of the FOs of E. floralis and B. africana above
the Niveau Goguel black shales (OAE 1a) is similar to the
order of FOs in the Cismon core of Italy (Fig. 5). Both
species occur above the ‘Livello Selli’ (Erba et al., 1999)
which is considered to be an equivalent of the Niveau
Goguel black shales (e.g., Coccioni et al., 1992; Bralower
et al., 1994). In contrast, Thierstein (1973) described the
FO of E. floralis in southeast France within the Niveau
Blanc, i.e., higher up in the succession. Black shale
horizons probably synchronous with the Niveau Goguel
have also been recognized in the Lower Saxony Basin
(Fischschiefer, Kemper & Zimmerle, 1978; Mutterlose,
1992; Bischoff & Mutterlose, 1998; Habermann &

Fig. 5. Composite lithological column of the Aptian–Lower Albian succession in the Vocontian Basin (SE France) with regional and supraregional
key beds plotted against biostratigraphy, and the distribution of Nannoconus spp. and Repagulum parvidentatum. Planktic foraminiferal and
ammonite biostratigraphy after Bréhéret (1997) and references therein), Moullade et al. (1998) and Kennedy et al. (2000). Right-hand side: first (FO)
and last (LO) occurrences of calcareous nannofossils. Arrow heads in black indicate major first and last occurrences (supraregional significance);
open arrow heads indicate minor first occurrences (regional significance). Nannoconid crises of the early Aptian and middle late Aptian are indicated
by NC I and NC II respectively. For lithological explanations see Fig. 3. D. des., Deshayesites deshayesi; T. bow., Tropaeum bowerbanki; P. nutfield.,
Parahoplithes nutfieldiensis; H. nolani, Hypacanthoplites nolani; L. tarde, Leymeriella tardefurcata; D. mammillatum, Douvilleiceras mammillatum; L.
cabri, Leupoldina cabri; G. ferreolensis, Globigerinelloides ferreolensis; G. alger., Globigerinelloides algerianus; H. trocoidea, Hedbergella trocoidea; C.
acutum, Corollithion acutum; C. litter., Chiastozygus litterarius; NG, Niveau Goguel; NB, Niveau Blanc; NC, Niveau Calcaire; NN, Niveau Noire;
NNC, Niveau Noire Calcaire; FA, Niveau Fallot; FN, Faisceau Nolan; FF, Faisceau Fromaget; DC, Délits Calcaire; NJ, Niveau Jacob;
NK, Niveau Kilian; HN, Haut Noire; NP, Niveau Paquier; LE, Niveau Leenhardt; L’Arb., L’Arboudeysse; P.-G., Pré-Guittard.
10 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22 11

Mutterlose, 1999) and known as the OAE 1a in the
Pacific and Atlantic oceans (e.g., Sliter, 1989; Bralower
et al., 1994).
The FOs of E. varolii and E. apertior were first
described from lower Aptian Tethyan sections where
both species occur in low numbers. In contrast, these
species are relatively common in sediments at high-
latitude locations (e.g., Bischoff & Mutterlose, 1998).
These species may have been overlooked previously or
referred to similar forms (group of Eprolithus) at lower
latitudes (compare Bown et al., 1998). The FO of
R. planus a little above the FO of E. floralis is similar to
the pattern described by Varol (1992) from different
Upper Aptian low-latitude sections.
The late Aptian part of the succession. The FOs of
Lapideacassis mariae, L. glans, Prediscosphaera sp. and
O. perspicuum occur in the lower Upper Aptian in the
Vocontian Basin (see Fig. 5). Hitherto, these taxa had
been recorded only from the uppermost Aptian
(Kennedy et al., 2000). Mutterlose (1996) recognized
Prediscosphaera spp. in the Tropaeum drewi (ammonite)
Zone (lower Upper Aptian) of the Lower Saxony Basin.
The FO of Prediscosphaera sp. observed within Niveau
Calcaire 2 in the Vocontian Basin (lower Upper Aptian)
seems to be similar in the Boreal and Tethyan sections.
The FO of Prediscosphaera columnata is situated 3 m
above the Délits Calcaire 2 (uppermost Aptian).
Thierstein (1973) described the FO of P. columnata from
within the Lower Albian in the upper part of the
Leymeriella tardefurcata (ammonite) Zone from the Col
de Palluel section in southeast France. Bréhéret (1997)
encountered the FO of P. columnata ‘just below DC2’.
Kennedy et al. (2000) recorded P. columnata 3 m above
the Niveau Jacob in the Pré-Guittard section. Above the
FO of P. columnata the FOs of Eiffellithus hancockii and
Cribrosphaerella ehrenbergii have been recorded.
The Niveau Jacob black shale is sandwiched between
the FOs of Prediscosphaera spinosa and P. columnata
(circular form), comprising an interval of 20 m. Based
on their two FOs, the biostratigraphy of the Niveau
Jacob is well constrained and can be correlated to black
shales in various regions of the world. Based on the FO
of P. columnata 2.5 m above it, the Livello 113 black
shale in Italy is synchronous with the Niveau Jacob in
the Vocontian Basin (Erba, 1988; Erba et al., 1989;
Bréhéret, 1997). Another black shale horizon rich in
ammonites occurs at the same stratigraphic level in the
North Atlantic at DSDP Site 401, and is interpreted as
an equivalent of the Niveau Jacob (Bréhéret, 1997).
The FOs of Helicolithus trabeculatus and Seri-
biscutum primitivum have been recorded below the
Niveau Kilian black shale. The FO of H. trabeculatus in
the Pré-Guittard section corresponds to observations
made by Kennedy et al. (2000) from the same section.
These authors, however, observed the FO of S. primi-
tivum at 10 m below the Niveau Leenhardt black shale
higher up in the Pré-Guittard section. Specimens of
H. trabeculatus are abundant in the interval studied. The
Niveau Kilian black shale possibly lies within the Monte
Nerone level in the Umbria-Marche Basin of Italy
(Erbacher, 1994).
The Aptian/Albian boundary. The Aptian/Albian
boundary is difficult to determine using marker species
(see Hart et al., 1996, Kennedy et al., 2000, and
Hancock, 2001 for detailed discussion). Recently,
Kennedy et al. (2000) defined the boundary on the basis
of the first occurrence of the ammonite Leymeriella
tardefurcata at the base of the Niveau Paquier black
shale in southeast France. In contrast, Hancock (2001)
suggested that the boundary would be better placed at
the base of the Lyelliceras lyelli Zone, which should be
recognizable on all continents. In this study, we have
followed Kennedy et al. (2000) and placed the Aptian/
Albian boundary at the base of the Niveau Paquier
black shale (see Fig. 5).
The lower Albian part of the succession. The FO and
LO of Arkhangelskiella stenostaurion (A. antecessor; see
Burnett, 1998) were recently described by Gale et al.
(1996) and Burnett (1997, 1998) from the upper Albian
Stoliczkaia dispar (ammonite) Zone of southeast France.
However, these authors studied only the Upper Albian
section and not the Upper Aptian. Our study indicates
that Arkhangelskiella stenostaurion makes up to 2% of the

Fig. 6. Nannofossils recorded. All illustrations in this figure and in Figs. 7 and 8 are light micrographs at a magnification of 3000 unless noted
otherwise. a–c Arkhangelskiella stenostaurion (Hill 1976) comb. nov.; XPL, Les Oustaus, Sample OUS 156; Lower Albian; b, as a, XPL; c, as a, PC.
d, e, Assipetra infracretacea (Thierstein, 1973) Roth 1973; XPL, Serre Chaitieu, Sample PSC 32; Lower Aptian; e, as d, PC. f, g, Biscutum constans
(Górka, 1957) Black, in Black & Barnes 1959; XPL, Tarendol, Sample LF 183/1/98; Upper Aptian; g, as f, PC. h, Braarudosphaera africana Stradner,
1961; XPL, Gaubert, Sample GAU 67; Upper Aptian. i, Braarudosphaera hockwoldensis Black 1972; XPL, Serre Chaitieu, Sample PSC 28; Lower
Aptian. j, Braarudosphaera regularis Black 1973; XPL, Serre Chaitieu, Sample SSC 1; Lower Aptian. k, Braarudosphaera sp.; XPL, Col de Palluel,
Sample LE 31; Lower Albian. l, Chiastozygus litterarius (Görka, 1957) Manivit 1971; XPL, l’Arboudeysse, Sample MP 50; Lower Albian. m,
Cribrosphaerella ehrenbergii (Arkhangelsky, 1912) Deflandre, in Piveteau, 1952; XPL, Col de Palluel; Sample LE 99, Lower Albian. n, o,
Diazomatolithus lehmanii Noël, 1965; XPL, Serre Chaitieu, Sample PSC 43; Upper Aptian; o, as n, PC. p, Discorhabdus ignotus Górka, 1957; XPL,
Col de Palluel, Sample LE 43; Lower Albian. q, Eiffellithus hancockii Burnett, 1998; XPL, Tarendol, Sample LF 161/2/98; Upper Aptian. r,
Eprolithus apertior Blacke 1973; XPL, Serre Chaitieu, Sample PSC 34; Lower Aptian. s, Eprolithus floralis (Stradner, 1962) Stover, 1966; XPL, Serre
Chaitieu, Sample SSC 21; Lower Aptian. t, Eprolithus floralis (Stradner, 1962) Stover, 1966; XPL, Serre Chaitieu, Sample PSC 35; Lower Aptian.
u, v, Eprolithus varolii Jakubowski, 1986; XPL, Serre Chaitieu, Sample PSC 43; Upper Aptian; v, as u, PC. w, Hayesites albiensis Manivit 1971;
XPL, l’Arboudeysse, Sample MP 65; Lower Albian. x, Hayesites albiensis Manivit, 1971; XPL, Les Oustaus, Sample OUS 138; Lower Albian.
12 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22 13

calcareous nannofossil assemblage within the Upper
Aptian Niveau Kilian black shale and is found frequently,
but in low numbers, in the Les Oustaus, l’Arboudeysse,
and Col de Palluel sections of the Lower Albian.
The FO of H. albiensis was observed below the
Niveau Paquier (46 m) in the Les Oustaus section.
Bralower et al. (1993) encountered the FO of H. albiensis
0.5 m below the Niveau Paquier black shale. They
focused, however, on the Niveau Paquier and did not
study the complete section from the Niveau Kilian to the
HN21 black shales. Within the interval from the Niveau
Kilian to HN21, Sollasites falklandensis first occurs
6.9 m below the Niveau Paquier. This observation cor-
responds to that of Bralower et al. (1993). The FO of
Placozygus cf. P. fibuliformis below the Niveau Paquier
black shale is similar to the record of Kennedy et al.
(2000) from the Pré-Guittard section, whereas the FO
of Kokia sp. has not been recorded from the latter,
probably because of the low numbers of this taxon.
The Niveau Paquier black shale is recognized in
several sections of the Tethyan region (Bréhéret, 1985,
1988, 1994; Arthur et al., 1990). It may correspond to
the Livello Urbino in Italy (Bralower et al., 1993) and
was recorded by Bréhéret (1985) from South Germany
near Hohenschwangau (‘In der Höll’) and from Austria
in the ‘Losenstein’ section. Several ODP/DSDP legs
from the North and South Atlantic contain probable
contemporaneous organic-rich sediments attributed to
the Oceanic Anoxic Event 1b (OAE 1b). These are
DSDP Site 545 on the Mazagan Plateau, ODP Site 641
on the Galicia Bank, DSDP Site 511 on the Falkland
Plateau, DSDP Site 386 on the Bermuda Ridge, and
ODP Site 1047 on the Blake Nose Plateau (Bralower
et al., 1993; Erbacher et al., 1999; Herrle, 2002).
6.2. Changes in calcareous nannofossil assemblages
The nannofloras studied from the Vocontian Basin
have a predominantly Tethyan (low-latitude) affinity in
the Aptian. This is indicated by the common occurrence
of species of Nannoconus which has been described as
a typical low-latitude taxon (e.g., Thierstein, 1976;
Mutterlose, 1989; Street & Bown, 2000). Its occurrence
in the Boreal Realm has been linked to warm surface-
water transgressions by Mutterlose (1989). The upper-
most Aptian–Lower Albian interval in the Vocontian
Basin is characterized by taxa more commonly associ-
ated with higher latitudes (Kennedy et al., 2000), such
as Repagulum parvidentatum and, in lower numbers,
Calculites anfractus and Seribiscutum primitivum.
Distribution of the boreal species Repagulum par-
videntatum. The nannofloral assemblages recovered
from the Aptian–Lower Albian interval in southeast
France show distinctive trends in their composition
which are probably of supraregional significance. The
abundances of the typical boreal species R. par-
videntatum (e.g., Mutterlose & Wise, 1990; Mutterlose,
1992; Street & Bown, 2000) are remarkable in a low-
latitude section (Fig. 5). Abundances of this species have
been recorded in high latitudinal settings in upper-
most Aptian–lower Albian deposits in the Wedell Sea
(Mutterlose & Wise, 1990; Mutterlose, 1992) and in the
North Sea area (Bown et al., 1998).
It is assumed that abundances of R. parvidentatum are
related to cool surface waters (Mutterlose, 1992; Erba
et al., 1992, Herrle, 2002). Therefore, at low latitudes
they may reflect a general cooling trend during the latest
Aptian–early Albian and an increased exchange of the
biota between the Boreal and Tethyan realms. A gener-
ally cool Aptian–early Albian interval was proposed by
Kemper (1987), based on the occurrence of glendonites.
This is in agreement with the oxygen isotopic compos-
ition of glendonites from dropstone facies in Australia,
which indicate cooler temperatures at high latitudes
during the late Aptian (De Lurio & Frakes, 1999;
Fig. 7). The assumption of cooling is also supported by
oxygen isotope variations based on benthic foraminifera
(Abreu et al., 1998).
The boreal influx of R. parvidentatum into the
Tethyan Realm was probably favoured by a long-term

Fig. 7. a, Helicolithus trabeculatus (Górka, 1957) Verbeek, 1977; XPL, Col de Palluel, Sample LE 131; Lower Albian. b, Lapideacassis glans Black,
1971; XPL, Col de Palluel, Sample LE 38; Lower Albian; c, as b PC. d, Lapideacassis glans Black, 1971; XPL, Serre Chaitieu, Sample SCH 129;
Upper Aptian. e, f, Lucianorhabdus salomonii Bergen, 1994; XPL, 2700; Serre Chaitieu, Sample SSC 31; Lower Aptian; f, as e, PC. g, Manivitella
pemmatoidea (Deflandre, in Manivit, 1965) Thierstein, 1971; XPL; 2700, Serre Chaitieu, Sample SC EAST 104; Upper Aptian. h, Micrantholithus
hoschulzii (Reinhardt, 1966) Thierstein, 1971; XPL, Serre Chaitieu, Sample SSC 5; Lower Aptian. i, Microstaurus chiastius (Worsley, 1971) Grün,
in Grün & Allemann, 1975; XPL, Serre Chaitieu, Sample NG 14; Lower Aptian. j, Nannoconus bucheri Brönnimann, 1955; XPL; 2700, Serre
Chaitieu, Sample PSC 12; Lower Aptian. k, Nannoconus circularis Deres & Achéritéguy, 1980; XPL; 2700, Serre Chaitieu, Sample PSC 12; Lower
Aptian. l, Nannoconus elongatus Brönnimann, 1955; XPL, Serre Chaitieu, Sample SSC 1; Lower Aptian. m, Nannoconus kamptneri Brönnimann,
1955; Brönnimann, 1955; XPL; 2700, Serre Chaitieu, Sample PSC 12; Lower Aptian. n, Nannoconus truittii Brönnimann, 1955; XPL, Serre
Chaitieu, Sample SC EAST 214; Upper Aptian. o, Nannoconus truittii Brönnimann, 1955; XPL, Serre Chaitieu, Sample PSC 12; Lower Aptian. p,
Nannoconus truittii Brönnimann, 1955; XPL, Serre Chaitieu, Sample PSC 9; Lower Aptian. q, Nannoconus vocontiensis Deres & Achéritéguy, 1980;
XPL; 2700, Serre Chaitieu, Sample PSC 28; Lower Aptian. r, Nannoconus wassallii Brönnimann, 1955; XPL, Serre Chaitieu, Sample SSC 1; Lower
Aptian. s, Owenia dispar (Varol, in Al-Rifaiy et al., 1990) Bown, in Kennedy et al., 2000; XPL, Serre Chaitieu, Sample PSC 19; Lower Aptian. t,
Owenia dispar (Varol, in Al-Rifaiy et al., 1990) Bown, in Kennedy et al., 2000; XPL, Serre Chaitieu, Sample PSC7; Lower Aptian. u, v Owenia
partitum (Varol, in Al-Rifaiy et al., 1990) Bown, in Kennedy et al., 2000; XPL, Serre Chaitieu, Sample PSC 28; Lower Aptian; v, as PC. w, Orastrum
perspicuum Varol, in Al-Rifaiy et al., 1990); XPL, Serre Chaitieu, Sample SC East 209; Upper Aptian. x, Orastrum perspicuum Varol, in Al-Rifaiy
et al., 1990; XPL, Serre Chaitieu, Sample SC East 209; Upper Aptian.
14 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22 15

sea-level rise (Haq et al., 1987), which improved the
exchange between low and high latitudes during the late
Aptian–early Albian (Figs. 5, 9 and 10). Moreover
ammonite faunas became more cosmopolitan at the
expense of Tethyan taxa during this period (Bréhéret &
Delamette, 1989).
Distribution of Nannoconus spp., A. infracretacea and
R. terebrodentarius. The up to 13-m-thick interval from
the Niveau Goguel (OAE 1a) to below the Niveau Blanc
is characterized by the absence of Nannoconus spp. (late
early Aptian NC6B nannofossil Subzone). A similar
absence of Nannoconus spp. has also been observed
within the OAE 1a of the La Bedoule succession in
southeast France (Moullade et al., 1998) and in Pacific,
Mexican, Italian and Spanish sections (Erba, 1994;
Aguado et al., 1999; Bralower et al., 1999; Luciani et al.,
2001). This interval corresponds to the nannoconid crisis
described by Erba (1994) and is here named nannoconid
crisis I (NCI; Figs. 5, 9 and 10).
In contrast to the distribution of Nannoconus
spp., the interval under discussion is characterized by
high numbers of large A. infracretacea (>6 µm) and R.
terebrodentarius (>10 µm). Common occurrences of
these two species in the upper Lower Aptian have also
been reported from low latitudes at DSDP Site 463
(Pacific), and from the Cismon core (Italy), the
Almadich Formation (SE Spain), and Mexican sections
(Erba, 1994; Aguado et al., 1999; Bralower et al., 1999;
Tremolada & Erba, 2002). Rucinolithus terebrodentarius
is also common within the lower Aptian Fischschiefer
in the Boreal Realm (North Sea Basin, Fischschiefer
Valhall V5; Jeremiah, 2001). In contrast A. infracretacea
and R. terebrodentarius are rare in the Upper Aptian–
Lower Albian succession of the Vocontian Basin and
then generally small. Owing to the restriction of large A.
infracretacea and R. terebrodentarius to the upper Lower
Aptian, they are interpreted as useful biostratigraphic
markers (Figs. 5 and 10).
Above the NCI, abundances of nannoconid taxa with
wide canals increase within the Niveau Blanc (N. globu-
lus, N. truittii, N. cricularis, N. bucheri, N. kamptneri;
Fig. 9). Similar observations have been made in Mexican
(Bralower et al., 1999) and Italian sections (Erba, 1994;
Cobianchi et al., 1997; Premoli Silva et al., 1999).
Above the Niveau Blanc in the lower Upper Aptian,
the abundance of nannoconids increases rapidly in the
middle part of the Globigerinellinoides algerianus
(foraminiferal) Zone, above the Niveau Fallot 4 black
shale. This section corresponds to the Nannoconus
truittii Acme described by Mutterlose (1989, 1991) an
‘event’ that has been observed in the Boreal as well as in
the Tethyan realms (Mutterlose, 1989; Erba, 1994).
A drastic decrease in Nannoconus spp. occurs at
the boundary of the Hedbergella trocoidea to Ticinella
bejaouaensis foraminiferal zones, which corresponds to
the lowest part of the Faisceau Nolan (FN1/FN2)
interval in the Vocontian Basin (Fig. 5). A similar rapid
decrease at the boundary between the H. trocoidea
and T. bejaouaensis zones is observed in Pacific and
Italian sections (Erba, 1994; Cobianchi et al., 1997). The
rapid decrease in Nannoconus spp. is, therefore, prob-
ably of supraregional significance and is named here
nannoconid crisis II (NCII; Figs. 5 and 10).
The middle to upper part of the Faisceau Nolan up to
the base of the Faisceau Fromaget is marked by moder-
ate abundances of nannoconids. Above the Faisceau
Fromaget, nannoconids occur in low numbers only. In
the Lower Albian, a rapid increase of nannoconids has
been recognized within the Niveau Paquier (OAE 1b;
Bréhéret et al., 1986), with percentages of up to 40%
(Herrle, 2002). OAE 1b black shales in Atlantic sections
and in Italy are characterized by the absence of nanno-
conids (Herrle, 2002). Therefore, the occurrence of
nannoconids within the Niveau Paquier is of regional
significance only.
Both nannoconid crisis I (prior to the OAE 1a) and
nannoconid crisis II are probably synchronous with
major drowning events of carbonate platforms (Fig. 10).
In general, nannoconids were one of the major
Tethyan marine carbonate producers during the Early
Cretaceous owing to their large size, compact structure,

Fig. 8. a, Palaeomicula maltica (Worsley, 1971) Varol & Jakubowski, 1989; XPL, Serre Chaitieu, Sample PSC 32; Lower Aptian. b, Polycostella?
prossii sp. nov.; XPL, l’Arboudeysse, Sample MP47; Lower Albian. c, Polycostella? prossii sp. nov.; XPL, Serre Chaitieu, Sample PSC 32; Lower
Aptian. d, Prediscosphaera columnata (Stover, 1966) Perch-Nielsen, 1984; XPL, Pré-Guittard, Sample PG 233/2/98; Upper Aptian. e, Predis-
cosphaera spinosa (Bramlette & Martini, 1964) Gartner, 1968; XPL, Les Oustaus, Sample OUS 79; Lower Albian. f, g, Radiolithus planus Stover,
1966; XPL, Serre Chaitieu, Sample SSC 24; Lower Aptian; g, as f, PC. h, Repagulum parvidentatum (Deflandre & Fert, 1954) Forchheimer, 1972;
XPL, Col de Palluel, Sample LE 131; Lower Albian. i, Repagulum parvidentatum (Deflandre & Fert, 1954) Forchheimer, 1972; XPL, Col de Palluel,
Sample LE 43; Lower Albian. j, Repagulum parvidentatum (Deflandre & Fert, 1954) Forchheimer, 1972; XPL, Tarendol, Sample LF 173/18; Upper
Aptian. k, Retecapsa tronickei sp. nov.; XPL, l’Arboudeysse, Sample MP47; Lower Albian; l, as k schematic view. m, Rhagodiscus cf. R.
achylostaurion; XPL, Col de Palluel, Sample LE 43; Lower Albian. n, Rhagodiscus achylostaurion; XPL, Col de Palluel, Sample LE 48; Lower
Albian. o, Rhagodiscus angustus (Stradner, 1963) Rheinhardt, 1971; XPL, Les Oustaus, Sample OUS 265; Lower Albian. p, Rucinolithus
terebrodentarius Applegate et al., in Covington & Wise, 1987; XPL, Serre Chaitieu, Sample PSC 8; Lower Aptian. q, Rucinolithus terebrodentarius
Applegate et al., in Covington & Wise, 1987; XPL, Serre Chaitieu, Sample PSC 9; Lower Aptian. r, Scapholithus fossilis Deflandre, in Deflandre &
Fert, 1954; XPL, l’Arboudeysse, Sample MP60; Lower Albian. s, t, Seribiscutum gaultensis Mutterlose, 1992; XPL, Les Oustaus, Sample OUS 99;
Lower Albian; t, as s, PC. u, Staurolithites mutterlosei Crux, 1989; XPL, l’Arboudeysse, Sample MP50; Lower Albian. v, Tranolithus minimus (Bukry,
1969) Perch-Nielsen, 1984; XPL, Col de Palluel, Sample LE 36; Lower Albian. w, x, Zeugrhabdotus erectus (Deflandre, in (Deflandre & Fert, 1954)
Reinhardt, 1965; XPL, Pré-Guittard, Sample PG 211/2/98; Upper Aptian; x, as w, PC.
16 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22

Fig. 9. Percentages of Nannoconus spp. and Assipetra infracretacea/Rucinolithus terebrodentarius in the lower 44 m of the Serre Chaitieu section
plotted against lithology and nannoplankton zones. Note the drastic increase in A. infracretacea/R. terebrodentarius and the low percentages in
Nannoconous spp. across the Niveau Goguel (OAE 1a) and the rapid increase of Nannoconus spp. within the Niveau Blanc. For lithological
explanations and abbreviations see Figs. 3 and 5.

and high abundances (e.g., Larson & Erba, 1999). Aptian possibly reflect a collapse in the marine carbon-
Therefore, the nannoconid crises and the platform ate system on a global scale (Herrle & Hemleben, 2001).
drowning of the late early Aptian and middle late We suggest that the observed nannoconid crises reflect
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22 17

Fig. 10. Compilation of the main calcareous nannofossil events against the magnetostratigraphy, absolute ages, planktic foraminiferal and ammonite
biostratigraphy, and the most important black shale and carbonaceous intervals in the Vocontian Basin. On the right the occurrence of nannoconids
and Repagulum parvidentatum, glendonites and platform drowning events, and a sea-level curve are plotted. Absolute ages compiled after Bralower
et al. (1997); planktic foraminiferal and ammonites zones after Bréhéret (1997) and references therein) and Kennedy et al. (2000); Barremian
stratigraphy after Moullade et al. (1998); glendonites after De Lurio & Frakes (1999); carbonate platform drowning according to Weissert et al.
(1998) and Lehmann et al. (2000); sea-level curve after Haq et al. (1987). For abbreviations see, Fig. 5.

the biotic response of a progressive increase in atmos- 7. Conclusions

pheric CO2 concentration owing to enhanced volcanic
degassing in the Aptian. The limiting effects of increas-
ing atmospheric CO2 to the calcite production of marine
calcifying phytoplankton species has recently been
shown by Riebesell et al. (2000). This view is also
supported by carbonate-platform drowning because a
rise in atmospheric CO2 levels causes a reduction in the
calcification rates in coral reefs (Langdon et al., 2000).
A detailed calcareous nannofossil biostratigraphy of
the Aptian–Lower Albian interval has been established
based on a composite section in the Vocontian Basin of
southeast France. It supports existing zonations, but
also provides a much higher resolution. It correlates well
with existing ammonite and planktic foraminiferal
zonation schemes for the vocontian basin.
18 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
The uppermost Aptian–Lower Albian is character-
ized by high abundances of R. parvidentatum, reflecting
a boreal influx into the Tethyan Realm. The high
abundances of this species at low latitudes probably
reflect a generally cool climate. This was accompanied
by a general rise in sea-level.
The distribution pattern of nannoconids is marked by
two significant crises: the late early Aptian (prior to the
Niveau Goguel, OAE 1a) nannoconid crisis I (NCI), and
middle late Aptian nannoconid crisis II (NCII). Both are
of supraregional significance and correlate with
carbonate-platform drowning events. Nannoconid crisis
I was followed by a proliferation of large A. infra-
cretacea and R. terebrodentarius. Both species are inter-
preted as important biostratigraphic marker species for
the upper Lower Aptian.
Acknowledgements
We thank Oliver Friedrich, Peter Kößler, and Jens
Lehmann for field assistance and André Bornemann
and Jörg Pross for valuable discussions and suggestions.
We are grateful to Christoph Hemleben, Hanspeter
Luterbacher and Volker Mosbrugger for access to their
facilities. Helpful comments of the reviewers Elisabetta
Erba and Paul Bown as well as from the editor, David
Batten, are highly appreciated. This research was sup-
ported within the framework of the collaborative
research centre SFB 275 (project A5) of the University
of Tübingen and by the German Research Foundation
(grant He 697/32-1/2).
Appendix A. Systematic palaeontology
Genus Polycostella Thierstein, 1971
Type species. Polycostella beckmanii Thierstein, 1971
Polycostella? prossii sp. nov.
Derivation of name. Named in honour of Dr Jörg
Pross, palynologist.
Holotype. Fig. 8b
Isotype. Fig. 8c
Type locality. Tarendol, southeast France, sample LF
181/2/98, uppermost Aptian.
Diagnosis (light microscope). Small nannolith, tenta-
tively assigned to the genus Polycostella because it
has 12 broad, radially organized, club-like elements
that are arranged symmetrically with respect to the
long and short axes and display a high birefringence.
Length 2.6–3.1 µm; width 2.8–3.3 µm.
Occurrence. Rhagodiscus angustus–Prediscosphaera
columnata calcareous nannofossil zones (Upper
Aptian), Serre Chaitieu, Tarendol (SE France).
Genus Retecapsa Black, 1971
Type species. Retecapsa angustiforata Black, 1971
Retecapsa tronickei sp. nov.
Derivation of name. Named in honour of Dr Jens
Tronicke, geophysicist.
Holotype. Fig. 8k, l
Type locality. L’Arboudeysse, southeast France,
sample MP47, Lower Albian
Diagnosis (light microscope). Medium-sized, nor-
mally elliptical placolith; species similar to R. schizo-
brachiata with respect to general construction and
light microscopic image but slightly smaller, and the
long axis of the ellipse is marked by a narrow bar,
whereas the short axis is characterized by four small
lateral bars. Under a light microscope the species is
moderately birefringent. Length 5.6 µm; width
4.3 µm.
Occurrence. Rhagodiscus angustus–Prediscosphaera
columnata calcareous nannofossil zones (Upper
Aptian–Lower Albian), Serre Chaitieu,
l’Arboudeysse (Col de Palluel) (SE France).
List of taxa recognized, with author attributions,
dates and notes
Calcareous nannofossils
Arkhangelskiella stenostaurion (Hill, 1976) comb.
nov.
1976 Broinsonia? stenostaurion Hill, pp. 127, 128,
pl. 3, figs 13–24
1996 Arkhangelskiella? sp. Burnett, in Gale et al.,
1996, fig. 4d
1998 Arkhangelskiella antecessor Burnett, p. 133
2000 Broinsonia? stenostaurion Bown, in Kennedy
et al., fig. 34u–y
Remarks. This species is referred to Arkhangelskiella
because of its rim structure, which is characterized by
two distal shield cycles with a darker outer edge, and
the distinctive central-area plate morphology. In con-
trast, the genus Broinsonia is restricted to forms with
broad central-area axial crosses and grills.
Assipetra infracretacea (Thierstein, 1973) Roth, 1973
Biscutum constans (Górka, 1957) Black, in Black &
Barnes 1959
Braarudosphaera africana Stradner, 1961
B. hockwoldensis Black, 1972
B. regularis Black, 1973
Braarudosphaera sp.
Calculites anfractus (Jakubowski, 1986) Varol &
Jakubowski, 1989
Chiastozygus litterarius (Górka, 1957) Manivit, 1971
Corollithion acutum Thierstein, in Roth & Thierstein,
1972
Cribrosphaerella ehrenbergii (Arkhangelsky, 1912)
 J.O. Herrle, J. Mutterlose / Cretaceous Research 24 (2003) 1–22
Deflandre, in Piveteau, 1952
Diazomatolithus lehmanii Noël, 1965
Discorhabdus ignotus Görka, 1957
Eiffellithus hancockii Burnett, 1998
Eprolithus apertior Black, 1973
E. floralis (Stradner, 1962) Stover, 1966
E. varolii Jakubowski, 1986
Hayesites albiensis Manivit, 1971
Helicolithus trabeculatus (Górka, 1957) Verbeek, 1977
Kokia sp. The taxon occurs very rarely in the Niveau
Paquier black shale.
Lapideacassis glans Black, 1971
L. mariae Black, 1971
Lithraphidites carniolensis Deflandre, 1963
Lucianorhabdus salomonii Bergen, 1994
Manivitella pemmatoidea (Deflandre, in Manivit,
1965) Thierstein, 1971
Micrantholithus hoschulzii (Reinhardt, 1966)
Thierstein, 1971
M. obtusus Stradner, 1963
Microstaurus chiastius (Worsley, 1971) Grün, in Grün
& Allemann, 1975
Nannoconus minutus Brönnimann, 1955
N. bucheri Brönnimann, 1955
N. circularis Deres & Achéritéguy, 1980
N. elongatus Brönnimann, 1955
N. globulus Brönnimann, 1955
N. kamptneri Brönnimann, 1955
N. quadriangulus Deflandre & Deflandre, 1967
N. truittii Brönnimann, 1955
N. vocontiensis Deres & Achéritéguy, 1980
N. wassallii Brönnimann, 1955
Nannoconus sp. 1 (wide central canal)
Nannoconus sp. 2 (narrow central canal)
Nannoconus sp.
Owenia dispar (Varol, in Al-Rifaiy et al., 1990) Bown,
in Kennedy et al., 2000
O. partitum (Varol, in Al-Rifaiy et al., 1990) Bown, in
Kennedy et al., 2000
Orastrum perspicuum Varol, in Al-Rifaiy et al., 1990
Palaeomicula maltica (Worsley, 1971) Varol &
Jakubowski, 1989
Placozygus cf. P. fibuliformis (Rheinhardt, 1964)
Hoffmann, 1970
Prediscosphaera columnata (Stover, 1966) Perch-
Nielsen, 1984
P. spinosa (Bramlette & Martini, 1964) Gartner, 1968
Prediscosphaera sp. The taxon occurs rarely in the
lower Upper Aptian of the sections studied.
Radiolithus planus Stover, 1966
Repagulum parvidentatum (Deflandre & Fert, 1954)
Forchheimer, 1972
Rhagodiscus achylostaurion (Hill, 1976) Doeven, 1983
Rhagodiscus cf. R. achylostaurion
Rhagodiscus angustus (Stradner, 1963) Reinhardt,
1971
19
R. asper (Stradner, 1963) Reinhardt, 1967
Rucinolithus terebrodentarius Applegate et al., in
Covington & Wise, 1987
Scapholithus fossilis Deflandre, in Deflandre & Fert,
1954
Seribiscutum gaultensis Mutterlose, 1992
S. primitivum (Thierstein, 1974) Filewicz et al., in
Wise & Wind, 1977
Sollasites falklandensis Filewicz et al., in Wind &
Wise, 1977
Staurolithites mutterlosei Crux, 1989
Tranolithus minimus (Bukry, 1969) Perch-Nielsen,
1984
Watznaueria barnesae (Black, in Black & Barnes,
1959) Perch-Nielsen, 1968
Zeugrhabdotus erectus (Deflandre, 1954) Reinhardt,
1965
Planktic foraminifera
Hedbergella trocoidea (Gandolfi, 1942)
Ticinella bejaouaensis (Sigal, 1960)
Ammonites
Hypacanthoplites jacobi Collet, 1907
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