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Research article

Journal of the Geological Society

Published Online First

doi:10.1144/jgs2015-150

Palaeocommunities, diversity and sea-level change from middle

Eocene shell beds of the Paris Basin
Stefano Dominici1* & Martin Zuschin2
1

Museo di Storia Naturale, Universit di Firenze, Via La Pira 4, 50121 Firenze, Italy
Department of Palaeontology, University of Vienna, Althanstrae 14, A-1090 Wien, Austria
* Correspondence: stefano.dominici@unifi.it

Abstract: The middle Eocene interval at some Paris Basin localities was studied through high-resolution stratigraphy.
Abundance data (499 species, 37 719 individuals) on the distribution of molluscs, collected at 12 shell beds of the middle
Lutetian and lower Bartonian, formed the basis for a palaeoecological study. The middle Lutetian succession is subdivided into
several elementary depositional sequences (EDS) interpreted as the product of relative sea-level change. Species-abundance
distributions are better correlated with EDS than with geographical locality, suggesting that sea level played an important role in
the distribution of palaeocommunities. Diversities were compared with analogous data from modern subtropical and warmtemperate intertidal and subtidal communities. We found that sea-level variation is responsible for a major change in the upper
part of the middle Lutetian succession, leading from high- to low-diversity palaeocommunities. From base to top sampled
palaeocommunities indicate a transition from high-energy and mesotrophic (EDS 2) to oligotrophic low-energy conditions of a
sandy lower shoreface (EDS 4) to an upper shoreface (EDS 5 and lower Bartonian), the last with mangroves and a seagrass
cover. Notwithstanding the Lutetian cooling, we found that subtropical conditions reached as far north as the Paris Basin. Our
study suggests that climatic fluctuations might be obscured by facies control.
Received 24 November 2015; revised 21 February 2016; accepted 9 March 2016

The middle Eocene of the Paris Basin belongs to one of the most
intensely studied stratigraphic intervals of geology. Shell beds there
have attracted scholars such as Guillaume Bruguiere, Jean-Baptiste
Lamarck, Grard Deshayes and Alcide dOrbigny, who contributed
to establishing the principles of modern stratigraphy and the tenets
of macroevolution (Rudwick 2005). As a corollary result of the
many efforts, the lists of its macro- and microfossils are mostly
completed (Merle 2008, and references therein; Courville et al.
2012; Lozouet 2014), but so far no species-abundance data of
quantitative bulk samples have been used to perform palaeocommunity and diversity analyses, apart from a recent study devoted to
one particular shell bed (Sanders et al. 2015). Such analyses,
however, are of major interest because fossil assemblages of the
Paris Basin show characters not known from benthic communities
living today at analogous latitudes, where many families and genera
have meanwhile undergone an almost complete turnover in
importance (Lozouet 2014; Tomasovch et al. 2014). For instance,
the Paris Basin was located around 3035N within the warmtemperate belt at a time of general cooling with respect to the Early
Eocene climatic optimum (EECO). Nonetheless, overall species
richness calculated at the stage level suggests values similar to those
of modern tropical hotspots (Huyghe et al. 2012a; Lozouet 2014;
Sanders et al. 2015).
The thickest part of the middle Lutetian succession is composed
of unlithified fine-grained sandstones, with a large component of
biogenic carbonates. Original aragonitic shells and residual colour
patterns are finely preserved and resemble much younger Neogene
shells (Merle 2008; Caze et al. 2011). Other well-studied
Palaeogene examples (e.g. the Pyrenees: Dominici & Kowalke
_
2014; SE Europe: Oppenheim 1901; the Middle East: Islamog
lu
et al. 2011; Harzhauser et al. 2012), whether siliciclastic or
carbonate-dominated, are poorly preserved. Similar good preservation is found only in some unlithified siliciclastic Bartonian
successions of the US Gulf Coast, including the Gosport Formation
(e.g. CoBabe & Allmon 1994) and the Stone City Formation

(e.g. Zuschin & Stanton 2002). Sedimentary structures, mudstone

partings and stratal surfaces are subtle, and massive bedding is
frequent, so that field correlations between Paris Basin outcrops
traditionally are not based on facies contrast but on macrofaunal
content (Gly 1996). The main bioclastic components are mollusc
shells, in concentrations of both sedimentological and biogenic
origin. Large foraminifera are also abundant, although their
accumulations do not take the form of the nummulitic banks that
are common in most coastal and shelfal Eocene successions of
nearby seas (Pyrenees: Dominici & Kowalke 2007; Huyghe et al.
2012b). If the Eocene is a strange old world (Clyde 1999), its
expression in the Paris Basin appears an utter mystery.
Paris Basin Palaeogene stratigraphy is relatively well understood,
with isopachous relations across long distances (Gly & Lorenz
1991; Gly 1996). This allows a good correlation of local
successions with regional subsidence patterns and global events.
Available studies recognize tectonic stability from the Danian to late
Bartonian (Robin et al. 2003). During this time, sedimentation was
controlled by major sea-level fluctuations, typically in the range of
third-order sea-level cycles (i.e. lasting 0.55 myr) and fourth-order
sea-level cycles, which are well framed in a chronostratigraphic
framework (Gly 1996; Huyghe et al. 2015). The third-order
Lutetian cycle, for example, was formed by three fourth-order
sea-level cycles. During the major transgression, which lasted more
than 3 myr (Huyghe et al. 2015), the coastline moved in a southern
direction in a stepwise fashion, with maximum flooding occurring
in the middle Lutetian, apparently without superimposed highfrequency fluctuations (Gly 1996).
To explore the ecological structure of middle Eocene shell beds
and shed further light on the history of sea-level changes, we have
reconsidered three classical Lutetian localities, well known at least
since the 18th century: Ferme-de-lOrme and Grignon, both near
Paris, and Fleury-la-Riviere, near Reims (Merle 2008; Courville
et al. 2012). In addition to the Lutetian localities, we included the
lower Bartonian succession at La Guepelle, to the NE of Paris, also

2016 The Author(s). Published by The Geological Society of London. All rights reserved. For permissions: http://www.geolsoc.org.uk/permissions.
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S. Dominici & M. Zuschin

one of the type areas for the definition of this stage (Pomerol 1981;
Fig. 1). The significance of sedimentary facies was reconsidered
and, to explore the nature of the Paris Basin biota, fossil
assemblages were bulk-sampled in some key units. Abundance
data on the distribution of molluscan species were evaluated and
used for a multivariate analysis and to compute sample-level
diversity. This allowed us to carry out for the first time a
palaeocommunity analysis of the Paris Basin marine macrofauna,
which provides information on the trophic structure of the benthic
ecosystem. A key question is whether the diversity of the Paris Basin
samples is comparable with that of modern tropical or warmtemperate environments. The sample diversity of the fossil
assemblages, also studied for the first time in the Paris Basin, was
therefore compared with that of modern assemblages from two wellstudied shallow marine settings, the northern Adriatic Sea and the
northern Red Sea, in warm-temperate and tropical climates,
respectively (Zuschin & Hohenegger 1998; Sawyer & Zuschin
2010). Species-level abundance data and the sample-level quantitative comparison opened a new approach to interpreting Paris Basin
shell beds. The influence of sequence architecture is also of major
interest because most changes in first and last occurrences of
species, and widespread changes in species abundance and
biofacies, occur at sequence boundaries and at major transgressive
surfaces (Holland 2000). This calls for evaluation of changes in
taxonomic diversity in the context of environmental bias to ensure
that these changes are not simply driven by sequence architecture
(Smith 2001).

Specifically, this approach allowed us to reconstruct Paris Basin

benthic palaeocommunities at a time of gradual change during the
middle Eocene and through climatic thresholds. The joint input from
sedimentary facies analysis and palaeoecology sheds light on external
factors controlling the record. This study allows a better understanding
to be obtained of the effects of global sea-level fluctuations in driving
both the structure and the composition of benthic communities.

Sedimentary facies
Paris Basin strata are roughly horizontal and isopachous (Gly
1996; Merle 2008), connecting outcrops as far as 160 km from one
another. Sections are oriented along depositional strike, which is the
reason for little lateral facies change within the studied succession
(Fig. 1). The most detailed sedimentological description of the
Faluniere de Grignon, where the Middle Lutetian is thickest and
most complete, has been given by Jean-Pierre Gly and Didier
Merle (cited by Guernet et al. 2012; Sanders et al. 2015). Those
researchers described and numbered the sedimentary units from 1 to
8, starting from the top (i.e. lower numbers represent younger units),
a basic framework adopted here. In this phase of our research,
middle Lutetian palaeoenvironments are interpreted based on recent
palaeontological studies carried out at Grignon (Guernet et al. 2012;
Huyghe et al. 2012a) and Fleury-la-Riviere (Courville et al. 2012;
see also Tomasovch et al. 2014) and from our own fieldwork.
Starting from the oldest, the sedimentary units of Gely & Merle
(cited by Guernet et al. 2012) are summarized as follows.

Fig. 1. Middle Lutetian at Ferme-de-lOrme, Grignon and Fleury-la-Riviere, and lower Bartonian at La Guepelle, in the Paris Basin, with position of bulk
samples. Small numbers and arrows at Grignon represent the extent of sedimentary units as defined by Guernet et al. (2012). Large numbers in circles
represent fifth-order depositional sequences as recognized in this paper, with main transgressive surfaces correlated according to the literature (Gly 1996;
Merle 2008). Description of the succession at La Guepelle is from B. Pattedoie ( pers. comm., 2007). A major unconformity separates the middle Lutetian
from the lower Bartonian (Huyghe et al. 2015; not all intervening strata are shown here). Insets show explanation of symbols and location map of the study
sites. Labels in rectangles refer to bulk samples collected in main shell beds.

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Paris Basin Middle Eocene shell beds

Units 7 and 8. Bioturbated calcareous sandstone with abundant
glauconite and quartz and sparse shell material. Thickness 34 m.
Based on field evidence, the association is characterized by
Cardium (Orthocardium) subporulosum dOrbigny 1850. A
tabular shell bed in the upper part is marked by sparse gravels and
Turritella shell debris. Fine gravels in the lower part of the Fleury
section, within sedimentological shell concentrations (Fig. 2a)
associated with bones of marine vertebrates, are here correlated with
the preceding shell bed. Units 7 and 8 have a complex stratigraphy,
at least in their lower part at Grignon. Samples PB27 and PB16 were
collected in the lower part of the Fleury-la-Riviere section, in
deposits correlative with the upper part of unit 7 at Grignon (Merle
2008; Courville et al. 2012).
Unit 6. Cross-bedded, locally bioturbated calcareous sandstones,
rich in quartz and glauconite, about 2 m thick. No samples were
evaluated from this unit.
Unit 5. Highly bioturbated massive calcareous sandstone with
abundant quartz and glauconite, about 1.5 m thick. In the lower part of
the unit, loosely packed shell beds are characterized by Campanilopa
giganteum (Lamarck 1804), which dominates the local biogenic shell
concentration with its high biomass (Fig. 2b). The upper shell beds,
with a species-rich mollusc association, are instead dominated by
Sigmesalia intermedia (Deshayes 1832) at Fleury-la-Riviere
(Courville et al. 2012). Sample PB9 was collected in unit 5 at Grignon.
Unit 4. Massive calcareous sandstone with shells in loosely
(Fig. 2c) or densely packed shell beds (Fig. 2d), or also resting on
irregular erosional surfaces in the basal part of the unit, in event beds
of sedimentological origin. Glauconite is rare. Thickness is about
2 m. The association is characterized in the field by Chama
lamellosa Lamarck 1806 and abundant Orbitolites complanatus
Lamarck 1801. Sample PB6 was collected at Grignon.

Units 2 and 3. Lithified massive calcareous sandstone about

2.5 m thick with very rare glauconite, sparse shell beds and
abundant Orbitolites in the lower part. In the upper part, the
association is dominated by Seraphs sopitus (Solander cited by
Brander 1776) and Avicularium lithocardium (Linn 1771), and
some colonial corals occur (Fig. 2f ). Samples PB1 and PB3 were
collected at Grignon, and PB10 at Ferme-de-lOrme.
Unit 1. Massive calcareous sandstone, about 0.5 m thick, with
thin laterally continuous shell beds (Fig. 2e, sample PB14). The
mollusc association is dominated in the field by Potamides
lapidorum (Lamarck 1804), Vicinocerithium echidnoides
(Lamarck 1804) and Saxolucina saxorum (Lamarck 1806), and
miliolids dominate among the benthic foraminifera.
The higher content of glauconite in older units may reflect lower
sedimentation rates, but the decreasing quartz content in the upsection direction gives evidence of the contrary.
Our investigations of the lower Bartonian succession cropping
out at La Guepelle, also known as Guepelle Sandstone Fm and
Beauchamp Sandstone Fm (biozone NP16: Huyghe et al. 2015),
are based on an unpublished description by B. Pattedoie ( pers.
comm., 2007). The third-order sequence boundary at the base of the
Bartonian succession (Fig. 1; see Huyghe et al. 2015) was not
observed in the field. Sedimentary facies at La Guepelle include a
number of massive sandstone units a few metres thick. They are
characterized by a sparse or loosely packed bioclastic fabric and
thinner calcareous intervals in the middle part of the succession,
topped by an important transgressive surface marked by boreholes
of bivalves (Fig. 1). These characters led previous researchers to the
interpretation of a lower shoreface environment, comparable with
that sampled in the middle Lutetian (e.g. Huyghe et al. 2015). A
coarsening-upward facies sequence above the transgressive surface,

Fig. 2. Middle Eocene shell beds of the

Paris Basin. (a) Sedimentological shell
concetration at Fleury-la-Riviere, near an
unconformity separating the middle
Lutetian from the Ypresian (EDS 2). (b)
Campanilopa giganteum biogenic shell
bed in plan view (Fleury-la-Riviere, EDS
2). (c) Loosely packed bioclastic fabric at
Grignon (EDS 4): disarticulated bivalves
interspersed with shell hash. (d) Densely
packed bioclastic fabric (Grignon, EDS 4;
height of outcrop about 20 cm):
disarticulated and nested bivalve shells
concentrated in pockets. (e) Potamides
lapidorum shell bed at Ferme-de-lOrme
(EDS 5): laterally continuous shell bed
within shelly sands. (f ) Shell bed in plan
view, with colonial corals and bivalves
(upper part of EDS 4, Grignon). For scale:
pencil, 16 cm; coin, 1.5 cm.

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S. Dominici & M. Zuschin

topped by coarse-grained sandstones with cross-bedding, indicates
the passage to a high-energy upper shoreface seafloor. Otherwise,
as for the middle Lutetian, a better palaeoenvironmental reconstruction is expected from a palaeoecological analysis of sampled
shell beds.

Sequence stratigraphy
The Palaeogene Paris Basin infilling is subdivided into five largescale depositional sequences of million-year duration, bounded by
major unconformities (third-order sequences in the sense of Haq
et al. 1987) and formed by the stacking of smaller units (Gly &
Lorenz 1991; Huyghe et al. 2015). The Lutetian third-order
sequence, of about 8 myr duration, has been divided into three
fourth-order sequences: sequence V (comprising the Glauconie
grossire Fm, the Calcaire Nummulites laevigatus Fm and the
Calcaire grossier Fm), sequence VI (Marnes et caillasses infrieures
Fm and Falun de Foulangues Fm) and sequence VII (Marnes et
caillasses suprieures Fm). These units approximately correspond to
sequences A, B and C of Gly (1996). Sequence V is of long
duration (about 5 myr) and comprises one sea-level fall dated
around the middle of biozone NP15 of calcareous nannoplankton
biostratigraphy, within Polarity Chron C20, followed by a new,
rapid transgressive event and a long, stepwise fall, until reaching a
major lowstand at the boundary C20C19 (Kominz et al. 2008).
Evidence for one episode of these global fluctuations is provided by
the fourth-order sequence boundary at the top of NP15 (VVI
unconformity of Huyghe et al. 2015).
The middle Lutetian localities under study here belong to the
highstand systems tract (HST) of sequence V. Around Paris, these
sediments sharply overlie glauconitic strata of the lower Lutetian,
whereas furthest west, near Reims, they rest on lower Eocene
deposits (Gly 1996), indicating an important erosion preceding the
major middle Lutetian transgression (Chron C20). The middle
Lutetian is formed by elementary units A6A10, interpreted as
parasequences (Gly 1996; Huyghe et al. 2012a). As an alternative
hypothesis, we propose that beds and bedsets are instead stacked to
form small-scale sequences bounded by subtle unconformities,
expressing low-amplitude fifth-order cycles and forming the
building blocks of the whole succession (elementary depositional
sequences: EDSs, numbered 16) (Fig. 1). Analogous highfrequency fluctuations, similarly expressed by metres-thick units,
are recognized in Lutetian carbonates of the Spanish Pyrenees,
increasing in amplitude and thickness in the Bartonian (Huyghe
et al. 2012b).
Facies changes justifying the grouping of bedsets into EDSs
include the following: (1) the presence of fine-grained gravels
(lower part of EDS 2 at Grignon and Fleury: Fig. 2a); (2) sharpbased coarse-grained strata with sedimentological shell concentrations (lower part of EDS 3 at Grignon and Fleury; lower part of EDS
4 at Grignon); (3) enrichment in carbonates in the middle part of
EDS 4; (4) occurrence of a shallow subtidal fauna in EDS 5. EDSs
24 are stacked to form a retrogradational pattern, with calcareous
sands of EDS 4 possibly representing a relatively deeper setting,
although not exceeding the lower limit of influence of wave action
during storms (i.e. lower shoreface). All 10 middle Lutetian bulk
samples belong to biozone NP15 of calcareous nannoplankton
stratigraphy (Huyghe et al. 2015). Samples were collected in the
transgressive tract of EDSs 25 (EDS 1 and EDS 6 were not
sampled), in biogenic shell beds from upper or lower shoreface
deposits (corresponding to nearshore and inner shelf samples,
respectively, of Tomasovch et al. 2014).
A superficial comparison suggests that also the Bartonian at La
Guepelle may be subdivided in a number of EDSs. Two sequence
boundaries are therefore proposed, with some transgressive surfaces
intervening in the succession (Fig. 1). Two bulk samples were

collected in the lower EDSs, which were possibly deposited in an

upper shoreface environment.
Most of the 12 samples were collected in a shell-rich and
bioturbated, massive sandstone lithofacies, indicative of low-energy
subtidal conditions below fair-weather wave base. Shallower
conditions were inferred from the presence of dispersed fine
gravels (PB27), abundance of intertidal gastropods (Potamides:
PB14), and position in a coarsening-upward succession (PBN7,
PBG). High glauconite content indicates low sedimentation rates.
Samples PB1 and PB3 were collected in a calcareous lithofacies
with denser accumulation of biogenic shell beds (Fig. 2d),
suggesting a further lowering of sedimentary input with respect to
underlying and overlying strata.

Analytical methods
Bulk samples (0.5 l) were washed through a series of sieves (1.0, 2.0
and 4 mm mesh-sizes). Species were separated and identified using
the monograph of Cossmann & Pissarro (19041913) and by
comparison with museum collections hosted at the University of
Florence, following modern taxonomic updates (Merle 2008). All
recognizable mollusc specimens in the fractions were counted and
combined for the purpose of this study. Raw abundances per
sample, for a total of 37 719 specimens and 499 species distributed
in 12 samples, and data on locality and stratigraphic interval served
as the basis for a Q-mode multivariate analysis. Cluster analyses
(Wards method) and ordination techniques (non-metric dimensional scaling, NMDS) and analysis of similarity (ANOSIM) were
applied to a matrix of square-root transformed per cent abundance
data. The similarity matrix for NMDS and ANOSIM is based on the
BrayCurtis Index. NMDS results are based on a 2D plot (stress =
0.1553), with Axis 1 explaining 47.42% and Axis 2 20.37% of the
variance. In the 3D plot, stress lowers to 0.115, with Axis 3
explaining only 7.52% of variance. In particular, ANOSIM
provides a way to test statistically whether there is a significant
difference between two or more groups of sampling units, based on
stratigraphy or geography. The R-values of the ANOSIM, which can
vary between zero (no difference between samples) and unity, were
used to evaluate relations between samples. Once groupings among
samples, recognized by cluster analysis and NMDS, were
established, the fossil fauna was used to explore the nature of cooccurrences by way of an R-mode analysis, carried out on a
simplified version of the dataset. This was obtained by including
only those species that contributed at least 2% of the total abundance
of at least one sample, and the species that co-occurred in at least
four samples, no matter what their frequency. By doing so, the
species list was downgraded to less than one-fifth of the original list,
and the total abundance decreased to just two-thirds (S = 91, n = 23
592). Results were expressed as frequency of a given species in the
original dataset. Per-sample, species-level diversity was measured
on the original dataset (S = 499, n = 37 719) through rarefied
species richness and evenness. Richness was calculated with 95%
confidence intervals at two values: n = 79, corresponding to the
minimum number of individuals (in sample PB3), and n = 623,
corresponding to the second lowest number of individuals (sample
94-6, from the Red Sea). Evenness was measured through the
ShannonWiener index. These values were compared with those of
modern analogues from the warm-temperate Adriatic Sea (Sawyer &
Zuschin 2010) and the subtropical Red Sea (Zuschin & Hohenegger
1998), which are taxonomically and taphonomically comparable
(i.e. the fauna consists mostly of molluscs) and were sieved on 1 mm
mesh-size. Mean values for the Red Sea, Adriatic Sea and Paris
Basin were measured for both values of richness and evenness. This
allowed us to evaluate the climate affiliation of Paris Basin shallow
marine biota, which during the middle Eocene inhabited latitudes
today associated with cool-temperate waters. All statistical analyses

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Paris Basin Middle Eocene shell beds

were carried out with the software package Primer 6.0 (Clarke &
Warwick 2001) and PAST 3.10 (Hammer et al. 2001)

Results
Multivariate statistics

Fig. 3. Multivariate analysis. (a) Cluster analysis. (b) Plot of NMDS;

samples grouped by clusters shown in (a). (c) Plot of NMDS; samples
grouped by locality. (d) Plot of NMDS; samples grouped by
chronostratigraphic unit (EDS).

In the Q-mode cluster analysis, two groups are recognized at a

similarity level of 15% (Fig. 3a). One larger group comprises all
middle Lutetian samples (cluster be), excluding PB14, the
youngest Lutetian sample in our analysis, which grouped with the
lower Bartonian sample PBG (cluster a). The second sample from
the lower Bartonian, PBN7, stands as an outlier. Within the middle
Lutetian cluster, three samples from EDS 4 at Grignon form a
smaller and tighter subset (cluster c: similarity above 40%). The
other three clusters within the large Lutetian group consist of two
samples each, which are always from different EDSs and different
localities (clusters ce).
The dataset includes four localities and five EDSs of chronostratigraphic significance, four from the middle Lutetian, at c.
42 48 Ma, and one from the lower Bartonian, at 3940 Ma. The
three samples from Fleury-la-Riviere, all collected within a 5 m
thick sandstone succession and located as much as 160 km from
the other section, are scattered in three groups. This suggests that
geographical position played a minor role in structuring the
distribution of the Paris Basin mollusc biota. Similarly, the two
samples collected at Ferme-de-lOrme, a section also only a few
metres thick, are also widely separated in the sample ordination plot
(Fig. 3c). The Q-mode clusters make more sense when interpreted in
terms of chronostratigraphic units. In the NMDS ordination, all EDS
4 samples, although from two different localities, plot close together
(Fig. 3d), with no overlap with those from other stratigraphic units.
EDS 2 and EDS 3 can be similarly separated. Displaced from these
are the sample from EDS 5 and the lower Bartonian samples. PB14,
from the middle Lutetian of Ferme-de-lOrme, plots relatively far
from the other middle Lutetian samples, and has affinities with the
lower Bartonian sample PBG as found in the cluster analysis
(Fig. 3a). This result is mirrored by the position of PB14 and PBG at
high values along Axis 1 of the NMDS plot (Fig. 3d). The ANOSIM
shows a lower Global R when the samples are grouped by
locality (Global R = 0.626; Table 1), than by EDS (Global R =
0.733; Table 2).
The R-mode cluster analysis highlighted eight main groups of
species contributing to the similarities of samples within each
EDS (Fig. 4, clusters labelled AH). Group A is the largest and
comprises 41 species, which preferentially occur in samples
collected from EDS 4 at Grignon and Ferme-de-lOrme. The
bivalve Paraglans calcitrapoides is abundant and, together with
the gastropods Vermicularia conica and Collonia striatus,
ubiquitous and exclusive of this interval. Cluster D also mainly
contributed to the grouping of samples from EDS 4. Some of the
species of clusters A and D, although not ubiquitous, were found
only in samples from this chronostratigraphic interval, including
Striarca decipiens, S. quadrilatera, Barbatia barbatula,
Cyclocardia squamosa, Plagiocardium granulosum, Emarginula
costata, Lapparentia fischeri and, in cluster D, Hipponix
cornucopiae, Sigmesalia trochoides and Siphonaliopsis minuta.
Species groups A and D, even when broken down to smaller
clusters, are composed of a mixture of trophic groups. These
comprise many suspension-feeding, shallow infaunal bivalves
(Carditidae, Cardiidae, Veneridae, etc.), many infaunal and
epifaunal carnivore gastropods (Natica, Vexillum, Crassispira,
etc.), some microherbivores (Tricolia, Pusillina) and parasites
(Tenuiscala). Cemented forms are rarer than in other clusters.
Among the species of cluster D, Eopleurotoma plicaria, Natica
epiglottina and Haustator mitis are particularly characteristic of
sample PB3 and also occur in sample PB16, from EDS 2; this
explains in part the grouping of these two samples in Q-mode
analyses (Fig. 3).

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S. Dominici & M. Zuschin

Table 1. Results of analysis of similarity (ANOSIM), factor locality
Groups
Grignon, Ferme
Grignon, Fleury
Grignon, Guepelle
Ferme, Fleury
Ferme, Guepelle
Fleury, Guepelle

R statistic

P-value

Possible permutations

Actual permutations

Number observed

0.491
0.508
0.982
0.5
0.75
0.917

0.095
0.018
0.048
0.1
0.333
0.1

21
56
21
10
3
10

21
56
21
10
3
10

2
1
1
1
1
1

Sample statistics (Global R): 0.626. Number of permutations: 999 (random sample from 83 160).

Table 2. Results of analysis of similarity (ANOSIM), factor EDS

Groups
EDS 4, EDS 3
EDS 4, EDS 5
EDS 4, EDS 2
EDS 4, Lower Bartonian
EDS 3, EDS 5
EDS 3, EDS 2
EDS 3, Lower Bartonian
EDS 5, EDS 2
EDS 5, Lower Bartonian
EDS 2, Lower Bartonian

R statistic

P-value

Possible permutations

Actual permutations

Number observed

0.600
1.000
0.836
1.000
0.000
0.750
0.500
1.000
0.000
1.000

0.095
0.167
0.048
0.048
0.667
1.000
0.333
0.333
0.667
0.333

21
6
21
21
3
3
3
3
3
3

21
6
21
21
3
3
3
3
3
3

2
1
1
1
2
3
1
1
2
1

Sample statistics (Global R): 0.733. Number of permutations: 999 (random sample from 83 160).

Cluster B comprises species occurring mostly in a sample from

EDS 3 (PB9, from Fleury-la-Riviere), such as suspension-feeding
(Haustator
imbricatarius)
and
carnivorous
gastropods
(Olivancillaria mitreola). Glycymeris pulvinata (a suspension
feeder) and Fustiaria circinata (a deposit feeder) are ubiquitous in
older Lutetian samples, disappearing up-section (EDS 5,
Bartonian). Overall, the suspension-feeding bivalves form the best
represented trophic group.
Clusters C and F contain species particularly abundant in EDS 2,
none of them being exclusive to this interval. Species belong to a
variety of trophic niches, with abundant microherbivorous gastropods Bittium semigranosum and B. transenna and many suspension-feeding bivalves and gastropods (e.g. Cardiidae, Veneridae,
Corbulidae, Turritellidae). Carnivores are less diversified or
abundant than elsewhere.
Clusters E and H contain species abundant in lower Bartonian
shell beds, the first with species of sample PBN7, the second with
species of PBG. Cluster H, characterized by the herbivorous
gastropods Potamides lapidorum and an unidentified rissoid, is
exclusive of younger samples (comprising EDS 5). Cluster E, in
turn, contains shallow infaunal suspension-feeding bivalves such as
Callocardia nitidula and Bicorbula gallica, and chemosymbiotic
lucinids (Parvilucina pusilla) that range from EDS 2 to the lower
Bartonian. Cubitostrea cubitus (cluster E) is found in both lower
Bartonian samples, whereas it is absent in the Lutetian. Finally,
cluster G groups species characteristic of sample PB14 from EDS 5
at Ferme. Here, characteristic species point to a vegetated bottom,
with abundant herbivorous gastropods Bittium duchasteli and
Tympanotonos semicoronatus. The chemosymbiotic lucinid
bivalve Parvilucina turgidula from cluster G points to organicrich bottoms with local sulphuric conditions, possibly associated
with the presence of seagrass.
A few species were found throughout the study interval, but the
majority are either middle Lutetian or lower Bartonian. Potamides
lapidorum is abundant and exclusive of EDS 5 and lower Bartonian
samples, whereas no species in the restricted dataset (S = 91) is
exclusive of the lower part of the middle Lutetian (EDS 2 or EDS 3).

Richness and evenness comparisons

Owing to differences in taxonomic and taphonomic conditions and
sampling design, there are not many modern examples that can be
compared with our fossil dataset. The two case histories that we
have selected are taxonomically and taphonomically comparable
with the Paris Basin study because they both consider the whole
benthic molluscan shelly fauna, including aragonitic shells, which
is well preserved in the Paris Basin collections. Sampling designs
are also comparable, with bulk samples sieved through the same
sieve sizes across the three studies that are being compared. The two
modern settings examined, the Red Sea and the Adriatic, like
the Paris Basin are from vast shallow seas enclosed by land masses.
The wide range of habitats included in both modern settings allows
for a meaningful comparison with the middle Eocene subtidal fossil
assemblage. In the end, we expect differences in composition,
evenness and abundance of single samples to be under the control of
climate, with the Red Sea and the Adriatic serving as templates for
subtropical and temperate settings, respectively. To facilitate
comparison, the information on bottom facies and water depth for
all samples is provided on a per-sample basis (Table 3).
The analysis confirms the uniqueness of samples from EDS 4:
this unit has the highest values of rarified species richness and
evenness among all samples, most notably PB1 and PB10, collected
in calcareous sandstones at Grignon and Ferme-de-lOrme,
respectively. The lowest evenness is present in lower Bartonian
samples at La Guepelle (PBN7 and PBG) and the upper part of the
middle Lutetian at Ferme-de-lOrme (PB14). Another sample with
low evenness is PB16, from the upper part of EDS 2 at Ferme. Our
diversity study therefore confirms a stratigraphic trend already
indicated by multivariate analysis: low or intermediate evenness in
samples from EDS 2 and EDS 3, very high values in EDS 4, and a
drop of evenness in EDS 5 and in the lower Bartonian. When Paris
Basin rarefied data are compared with their modern analogues, we
find that values encountered in EDS 4 are similar to those measured
in assemblages from modern coral sand and reef slope habitats; that
is, possibly the highest diversities encountered in the tropical Red

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Paris Basin Middle Eocene shell beds

Fig. 4. R-mode cluster dendrogram (analysis conducted on a subset of the original dataset) compared with sample composition, in stratigraphic order, with
frequencies expressed semiquantitatively.

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S. Dominici & M. Zuschin

Table 3. Comparison of rarefied species richness and evenness of molluscan faunas from samples collected in the modern Red Sea (Zuschin & Hohenegger
1998), the modern Adriatic Sea (Sawyer & Zuschin 2010) and in the middle Eocene of the Paris Basin (this study)

Bottom facies

Water depth (m) Sample label

ShannonWiener
Index

Rarefied species richness

EDS

Richness (n = 79) 95% CI Richness (n = 623) 95% CI

Red Sea
Coral sand
Coral sand
Coral sand
Coral sand
Muddy sand
Muddy sand
Mud
Mud
Reef slope
Mangrove
Reef slope
Sandy seagrass
Muddy seagrass
Red Sea (mean)
Adriatic Sea
Mud
Mud
Mud
Delta sandy claysilt
Gravelly sand
Gravelly sand
Tidal flat
Tidal flat
Adriatic Sea (mean)
Paris Basin
Sand
Sand
Sand
Sand
Sand
Sand
Sand
Sand
Sand
Sand
Sand
Sand
Paris Basin (mean)

Shannon_H

10
10
10
10
23
23
39
39
19
<1
12
6
40

94-1-a
94-1-b
94-1-c
94-1-d
94-3-a
94-3-b
94-4-a
94-4-b
94-5
94-6
95-31
B-5-8
C-1-3

40.55
41.24
37.31
39.60
25.83
23.60
17.62
19.95
43.30
24.58
41.57
32.08
23.87
31.62

1.11
1.09
1.12
1.10
1.17
1.19
1.00
1.08
1.05
1.13
1.09
1.14
1.22

130.40
132.57
119.43
127.14
75.63
68.80
37.22
45.86
135.43
69.00
131.26
91.55
70.90
95.01

0.93
0.92
0.89
0.90
0.47
0.63
0.92
0.93
0.79

13
14
11
4
6
7
Intertidal
Intertidal

Panzano_2
Panzano_4
Panzano_6
Panzano_8
Panzano_0
Panzano_00
Wattkante
Boxcore

19.18
16.32
16.73
26.13
31.59
27.82
9.86
9.42
19.63

1.04
1.06
1.09
1.00
1.10
1.13
1.03
0.87

38.96
33.08
35.55
56.32
78.46
71.13
20.15
16.38
43.75

0.75
0.76
0.76
0.66
0.84
0.88
1.02
0.86

2.61
2.36
2.41
3.14
3.53
3.20
1.57
1.70
2.56

Upper shoreface
Upper shoreface
Upper shoreface
Lower shoreface
Lower shoreface
Lower shoreface
Lower shoreface
Lower shoreface
Lower shoreface
Lower shoreface
Lower shoreface
Upper shoreface

PBG
PBN7
PB14
PB4
PB10
PB1
PB3
PB6
PB9
PB22
PB16
PB27

13.51
17.88
21.53
26.23
41.33
43.95
36.00
33.13
22.12
21.96
20.56
23.49
29.86

0.94
1.01
1.03
1.17
1.00
0.99

23.33
32.95
43.79
67.64
96.75
109.13

0.72
0.41
0.66
0.65
0.70
0.74

1.06
1.11
1.17
0.99
1.13

82.10
53.12
55.31
37.58
61.80
70.43

0.75
0.78
0.91
0.63
0.61

2.26
2.52
2.82
2.80
4.04
4.21
3.39
3.62
2.93
2.79
2.63
2.94
3.26

Lower Bartonian
Lower Bartonian
EDS 5
EDS 4
EDS 4
EDS 4
EDS 4
EDS 4
EDS 3
EDS 3
EDS 2
EDS 2

0.96
0.93
1.07

4.12
4.15
3.90
4.04
2.91
2.75
2.65
2.83
4.18
2.81
4.04
3.44
2.84
3.44

Samples from EDS 2 to EDS 5 are from the middle Lutetian; samples PBG and PBN7 are from the lower Bartonian.

Sea (Table 3; Zuschin & Hohenegger 1998). In contrast, in EDS 5

and the lower Bartonian shell beds the palaeocommunities show the
high dominance typical of intertidal assemblages from the modern
warm-temperate Adriatic Sea (Table 3; Sawyer & Zuschin 2010).

highlights three possible end-members of a continuum, where factors

such as nutrient level, water depth and trophic interaction with plants
contribute to shaping macrofaunal benthic communities.

Mesotrophic sandy bottoms

Middle Eocene molluscan palaeoecology
The quantitative analysis of middle Eocene Paris Basin shell beds has
highlighted a change in the composition of mollusc palaeocommunities that parallels and expands knowledge based on lithology and
stratigraphy. Evenness and complexity in trophic relationships
generally increase from sandstones of EDS 23 to calcareous
sandstones of EDS 4, dropping to a minimum in calcareous
sandstones of EDS 5 and lower Bartonian sandstones. This change
is also reflected in the relative position of samples in ordination space
(Fig. 3), although relationships here are more complex to interpret and
a simple gradient cannot be extrapolated. The following discussion

The lowermost sample PB16, from the middle Lutetian (EDS 2),
and PBN7, from the lower Bartonian, show low evenness (Table 3),
but they occupy different positions in the NMDS plot (Fig. 3).
Dominant species of either assemblage, however, belong to the
suspension feeders. These include a turritelline gastropod,
Haustator mitis (32% of frequency in PB16), and two infaunal
venerid bivalves, Calloocardia nitidula and Callista elegans
(respectively 32% and 23% in PBN7). Suspension feeders can act
as opportunist species within shallow marine benthic communities,
taking advantage of high quantities of seston and capable of facing
rapid changes of environmental conditions. Accordingly, high

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Paris Basin Middle Eocene shell beds

dominance of turritelline gastropods is frequently associated with
deltaic and upwelling zones, where seasonally driven nutrients
abound (Allmon 1976). As upwelling is unlikely, given the
palaeogeographical setting of the Paris Basin, some form of
deltaic influence can be hypothesized for the PB16 palaeocommunity. In contrast, rapid burrowers such as the dominant venerid
bivalves in PBN7 are abundant in high-energy shallow subtidal
bottoms, where they can withstand the vagaries of a rapidly shifting
sandy substrate and take advantage of the abundant suspended
organic matter (Stanley 1977). The separation between PB16 and
PBN7 in the ordination plot can thus be explained by two different,
but similarly unpredictable subtidal environments. An important
species-level turnover has, however, taken place during the several
million years that separate the two. This calls for also considering
evolutionary change in explaining the distance of PBN7 from all
other samples. This is not the case for PB3, also from the middle
Lutetian (EDS 4), which shares with PB16 abundant Haustator
mitis (eight individuals, 32% in PB16), explaining their proximity
in ordination space. Rarefied species richness and evenness of this
sample are relatively high, but have to be interpreted with caution
because of the small size of the sample (n = 79).

Oligotrophic sandy bottoms

All other middle Lutetian samples are plotted in the middle part of
the NMDS. Samples from EDS4, including PB1, PB4 and PB6
from Grignon and PB10 from Ferme-de-lOrme, have the highest
evenness, comparable with that of open marine coral sands of
subtropical settings such as the Red Sea (Table 3). Similarly, we can
assume that the upper half of EDS 4 was deposited under
oligotrophic conditions in an ecosystem characterized by narrow
niches. Accordingly, the palaeocommunity is evenly composed of
herbivores (Emarginula costata, Collonia stratus, Ptychocerithium
inabsolutum, Vicinocerithium sp., Tricolia sp., Pusillina nana,
Lepidochitona grignonensis), carnivores (Tenuiscala cloezi,
Payraudeautia perforata, Lapparentia fischeri, Volvarinella spp.,
Eopleurotoma plicaria, Cryptoconus spp.), some deposit feeders
(Rimella fissurella), and suspension feeders. The last are in turn
evenly represented by byssate epifauna (Barbatia barbatula,
B. angusta, Striarca decipiens), cemented epifauna (Vermicularia
conica, Vermetia sp.) and infauna (Paraglans calcitrapoides,
Crassatina triangularis, Varicorbula minuta, Plagiocardium
granulosum, etc.). These characters suggest a subtidal setting
deeper and furthest from terrestrial inputs with respect to all other
palaeoenvironments considered.
Among samples from EDS 4, PB10 shows many similarities to
PB22 from EDS 3, including the abundance of the herbivore Bittium
transenna, the suspension-feeding Turritellidae (Sigmesalia intermedia and Haustator funiculosus) and the Veneridae Callista
elegans and Cyclocardia serrulata. Because the last two are
abundant also in PBN7 from high-energy subtidal bottoms, and
because the Turritellidae more often prefer high-nutrient settings,
we infer that Ferme-de-lOrme and Fleury-la-Riviere, where PB10
and PB22 were collected, respectively, were shallower and closer to
terrestrial inputs with respect to Grignon. The last two remaining
assemblages to consider, PB27 and PB9 from EDS 2 and 3,
respectively, can also be interpreted as representing more
unpredictable palaeoenvironments than those encountered at
Grignon within EDS 4. Evenness is not as high as in the latter,
and opportunistic suspension feeders often abound at the expense of
other trophic groups; this is the case with Haustator funiculosus
(23% in PB27), Cubitostrea plicata (21% in PB9, 17% in PB27)
and Glycymeris pulvinatus (20% in PB9, 2% in PB27).
Analogies of Paris Basin rarefied diversities with those measured
in modern shallow marine mollusc communities in the Red Sea
strengthen the above palaeoecological interpretation. Modern coral

sands are good analogues of samples PB1, PB4, PB6 and PB10
(EDS 4, excluding PB3) because they have even proportions of the
main trophic groups (herbivores, suspension feeders and carnivores), and because opportunistic suspension feeders such as
Turritella or Corbula are missing (Zuschin & Hohenegger 1998).
The lack of chemosymbiotic bivalves in EDS 4 palaeocommunities,
although they are diversified and abundant in Red Sea subtidal coral
sands, can be explained by the lack of a seagrass cover in Paris Basin
bottoms during this part of the middle Lutetian.

Mangroves and seagrasses

Younger samples PBG and PB14 occupy the lower right quarter of
the ordination, forming a loose cluster (Fig. 3a). Their trophic
composition justifies, for both, the interpretation of palaeocommunities from a very shallow subtidal sandy bottom, with very high
organic content from the decomposition of vegetal matter. The
simplest palaeocommunity is represented by lower Bartonian
sample PBG with a high dominance of herbivorous gastropods of
the families Potamididae (Potamides lapidorum, Tympanotonos
semicoronatus), Cerithiidae (Ptychocerithium lamellosum) and
Rissoidae (Pseudotaphrus buccinalis and an undetermined
species), followed by the carnivores (Amauropsina canaliculata, a
naticid) and some suspension feeders, either byssate- (Trinacria
media) or cemented-epifaunal (Cubitostrea cubitus), with only rare
infauna (Cyclocardia pulchra).
Next in composition comes sample PB14 from the upper part of
the middle Lutetian (EDS 5). It shares with PBG the dominance of
cerithioidean herbivorous gastropods, mostly belonging to the same
species (Potamides lapidorum, Tympanotonos semicoronatus,
Ptychocerithium lamellosum), but also the cerithiids Bittium
duchasteli and B. semigranosum. Rissoidae, however, are absent;
their ecological role is played in part by a Pseudomelaniidae
(Bayania sp.), and trophic complexity and evenness are considerably higher than in PBG. In particular, the suspension feeders are
much more diverse and equally distributed within the epifauna (e.g.
Trinacria deltoidea, Striarca quadrilatera, Barbatia angusta) and
the infauna (e.g. Cyclocardia serrulata, Gari dutemplei, Venerella
secunda, Katelysia texta). Alongside the suspension feeders, the
infauna is occupied by abundant chemyosymbiotic lucinid bivalves
(Parvilucina turgidula), indicative of a substratum rich in organic
content. Carnivores are represented by rare Oliviidae
(Olivancillaria mitreola) and Marginellidae (Volvarinella
eburnea).
Modern Potamididae show parallel distribution with mangrove
trees along the tropical intertidal belt (for the relations between
Eocene and modern potamidids, see Reid et al. 2008; Dominici &
Kowalke 2014), but the presence of Rissoidae (PBG) and miliolid
foraminifera (PB14), together with lucinid bivalves, points also to
the presence of seagrass. The comparison of rarefied richness and
evenness for these two samples with those from the Red Sea points
to a species-abundance distribution similar to that of a modern
mangrove environment (Table 3). We therefore suggest a very
shallow subtidal environment, closely associated with a mangrove
ecosystem, possibly covered with sparse seagrass.

Discussion
Several aspects complicate a sequence stratigraphic interpretation:
the thinness of the sections (515 m) and the slow net sedimentation
rates they imply, the pervasive bioturbation that hampers the
development of a facies model, the paucity of outcrops and the
orientation of the section along depositional strike. However,
available data on the sequence architecture of low-accommodation
Cenozoic siliciclastic successions allow our sequence stratigraphic
interpretation to be considered as a viable working hypothesis. In

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S. Dominici & M. Zuschin

fact, data-rich studies from similar closed marine embayments
developed along passive continental margins allow for the
recognition of similarities. A very good analogue is the middle
Eocene Clairborne Group, in the onshore US part of the Gulf of
Mexico Basin (Hackley 2012). Claiborne formations contemporaneous with those here under study are the Welch Formation
(middle Lutetian), the Sparta Sand and the Cook Mountain
Formation (lower Bartonian). Similarly to their Paris Basin
analogues, they represent shallow marine sediments and are only
a very few metres thick, glauconitic, richly fossiliferous and laterally
persistent for tens of kilometres (Stenzel 1940; Gimbrede 1962;
Yancey & Davidoff 1994), implying low sedimentation rates and an
even topography. A younger analogue is found in the Miocene of
Maryland, further north along the US east coast. This is
characterized by laterally continuous and rich shell beds cropping
out for tens of kilometres in a strike direction, with unconformitybounded regressivetransgressive cycles stacked to form a retrogradation succession lasting more than 10 myr, but only about 30 m
thick (Kidwell et al. 2015).
The sedimentary facies analysis and palaeoecology of middle
Eocene Paris Basin shell beds allow an interpretation of the nature of
the shallow marine benthic ecosystem and its abiotic factors during
a peculiar time in the Cenozoic. The doubthouse comprised a
period between the Early Eocene greenhouse climate, with the
highest temperatures of the Cenozoic, and the Oligocene icehouse,
featuring a sharp decrease in global temperatures and the formation
of the Antarctic ice sheet. This period was characterized by a global
temperature decrease interrupted by a temporary amelioration
during the lower Bartonian (the Middle Eocene Climatic
Optimum; MECO) lasting about 500 kyr (Bohaty & Zachos
2003). Palaeoenvironmental change during the middle Lutetian
has been recently evaluated by geochemical and palaeontological
studies conducted on the Grignon succession, which are directly
comparable with our approach (Fig. 5). The isotopic study on
mollusc shells (Huyghe et al. 2012a, 2015) and the study on
ostracod assemblages (Guernet et al. 2012) suggest that maximum
depths were reached in the middle part of their parasequence A6,
corresponding to the lower part of EDS 2, in our study represented
by sample PB27 from Fleury-la-Riviere (Figs 1 and 5). In our
interpretation, however, PB27 is representative of a shallowing with
respect to underlying sediments (Glauconie grossiere Fm: Huyghe
et al. 2015; EDS 1) because of its palaeoecological signal (i.e.
relatively low evenness and high abundance of opportunistic
suspension feeders). Sedimentary facies analysis and the palaeoecology of mollusc shell beds suggest instead that the middle
Lutetian (fourth-order) maximum flooding surface (MFS) coincided with the lower part of shelly calcareous sandstones in the
upper part of the section, at the MFS (fifth-order) of EDS 4 (Fig. 5).
This interval is also marked by the richest ostracod assemblage
(Guernet et al. 2012). This fourth-order MFS would thus pinpoint
the beginning of the sea-level highstand recognized in global sealevel curves at around 45.0 Ma (Kominz et al. 2008). The sequence
boundary separating EDS 4 and EDS 5 would therefore mark the
long and stepwise sea-level drop occurring between about 44 and
43 Ma. This resulted in a major change in shell bed composition
recorded during the ensuing transgression, which is correlated with
the global rise occurring at about 42 Ma (Kominz et al. 2008), when
only very shallow subtidal depths were attained (sample PB14,
collected at Ferme-de-lOrme). A short-lived sea-level fluctuation is
recorded at Ferme-de-lOrme at the base of a thin muddy interval
( possibly deposited in an intertidal environment), overlain by upper
shoreface deposits dominated in the field by the turritellid
Sigmesalia (nutrient-rich shoreface environment; Fig. 1). The sea
level then fell again, in coincidence with the pre-MECO climatic
cooling. During this time, the third-order sequence boundary was
formed, separating the Lutetian from the Bartonian (Huyghe et al.

Fig. 5. Middle Lutetian stratigraphic units recognized in this study,

compared with units described by previous researchers. All middle
Lutetian samples used in our palaeoecological analysis were projected
onto the Grignon succession, taken as a template of this time interval.

2015). The La Guepelle succession can be correlated with the

general transgression occurring during the MECO, centred at
around 40.0 Ma (base of Chron C18n.2n: Huyghe et al. 2015).
Similarly to the transgressive systems tract of EDS 5 and EDS 6,
depths did not attain the same magnitude as in the middle Lutetian.
Cooling during the later part of the middle Lutetian, and the further
temperature drop preceding the MECO (Bohaty & Zachos 2003),
may explain the species-level turnover registered at the stage level
(Cossmann & Pissarro 19041913). Our sample-level data,
however, point to a facies shift at the sequence boundary between
EDS 4 and EDS 5, which controls mollusc diversity. We therefore
suggest that sequence stratigraphical architecture controls patterns
of faunal change at this regional scale, as has been shown for several
other regions and time intervals (e.g. Brett 1998; Patzkowsky &
Holland 1999; Smith et al. 2001; Zuschin et al. 2011).
The middle Eocene history of mollusc benthic communities
recorded in the Paris Basin is just one step in the long and
complex march leading from early Eocene greenhouse shallow
marine ecosystems to the modern fauna (Lozouet 2014). In this
time interval, the macroevolutionary turnover was particularly
severe at intertidal and shallow subtidal depths (Tomasovch et al.
2014). Middle Lutetian communities, however, are similar to
modern tropical assemblages such as those living in coral sands

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Paris Basin Middle Eocene shell beds

and seagrass bottoms of the Red Sea, sharing high evenness and
complex trophic relationships (Table 3; Zuschin & Hohenegger
1998), and unlike warm-temperate faunas of the Adriatic Sea,
which are dominated by fewer species at all depths (Table 3;
Sawyer & Zuschin 2010). This evidence confirms previous
suggestions (e.g. Huyghe et al. 2012a) that, even during the
middle Eocene cooling, subtropical conditions of the Tethyan
realm (Lozouet 2014) reached as far north as the Paris Basin. The
same situation was not found in the later part of the middle
Lutetian, nor during the MECO, notwithstanding an important
global climatic amelioration. Before concluding that shell beds of
EDS 4 record the last evidence of tropical affinity among midlatitude shallow marine communities in Europe, younger assemblages from a comparable facies must be sought.

Conclusions
Important middle Eocene successions of the Paris Basin were
studied, starting from a re-evaluation of sedimentary facies and
through new bulk-sampling in mollusc shells beds. The study
interval comprises the middle Lutetian and the lower Bartonian,
which are separated by a major depositional sequence boundary on a
regional scale. Our study reached the following conclusions.
(1) Both intervals can be subdivided into small-scale
unconformity-bounded units (i.e. metre-scale elementary
depositional sequences; EDSs), interpreted as the product of
high-frequency (fifth-order) cycles of sea-level change. We
identified five EDSs in the middle Lutetian and two in the
lower Bartonian. Sequence boundaries were recognized at the
base of sandstones with fine gravels or cross-bedding, within
successions dominated by bioturbated fine-grained shelly
sandstone and shelly calcareous sandstone.
(2) The species abundance distribution of molluscan assemblages
from 12 bulk samples of major shell beds, used for
multivariate analysis and diversity analysis, helped
reconstruct the composition and structure of the benthic
communities living at this peculiar time of Palaeogene climate
transition. Palaeocommunities from middle Lutetian EDS 14
are marked by high evenness and trophic complexity, which
peaked in EDS 4. These indicators dropped in EDS 5, possibly
in EDS 6 (not sampled), and in lower Bartonian shell beds.
(3) When compared with modern assemblages, middle Lutetian
communities strongly resemble mollusc species from coral
sands at subtropical latitudes, but less so warm-temperate
shallow marine communities. This is counterintuitive if Paris
Basin palaeolatitudes are considered. It does, however, agree
with previous knowledge on stage-level species richness,
calculated after more than two centuries of research on these
successions. We therefore confirm that the middle Lutetian
Paris Basin constitutes the remnant of a former early Eocene
greenhouse world, which disappeared as the icehouse world
of the EoceneOligocene boundary was approached.
(4) The steps with which the climatic transition occurred cannot
be understood unless the appropriate facies is sampled in
younger sediments. This was not possible at the study sites
because EDS 5, EDS 6 and Bartonian assemblages are from
shallower water depths. The vertical stacking of Paris Basin
palaeocommunities suggests that the peak of tropical diversity
of EDS 4 coincides with a fourth-order sea-level highstand
recorded on a worldwide basis.

Acknowledgements and Funding

The following people ensured the completion of this paper: D. Merle,
B. Pattedoie, P. Legrand, A. Tomasovch and M. Stachowitsch. We are grateful
to S. Holland for a thorough review of the paper. This work was supported by the

Austrian Science Fund (FWF) project P19013-Bio and MIUR/PRIN 20102011

(Past Excess CO2 worlds: biota responses to extreme warmth and ocean
acidification).

Scientific editing by Quentin Crowley

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