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Research article
doi:10.1144/jgs2015-150
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
2016 The Author(s). Published by The Geological Society of London. All rights reserved. For permissions: http://www.geolsoc.org.uk/permissions.
Publishing disclaimer: www.geolsoc.org.uk/pub_ethics
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.
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
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
Results
Multivariate statistics
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).
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).
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.
Bottom facies
ShannonWiener
Index
EDS
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.
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
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.
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
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.
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