Journal of Sedimentary Research, 2011, v.

81, 641655
Research Article
DOI: 10.2110/jsr.2011.53

MODERN HETEROZOAN CARBONATES FROM A EUTROPHIC TROPICAL SHELF (MAURITANIA)

JULIEN MICHEL,* GUILLEM MATEU VICENS,{

AND

HILDEGARD WESTPHAL{

MARUM and Department of Geosciences, Universitat Bremen, Leobener Strae, 28359 Bremen, Germany
e-mail: hildegard.westphal@zmt-bremen.de

ABSTRACT: Heterozoan or foramol production is typical in extratropical carbonate sedimentary systems. However, under
mesotrophic to eutrophic conditions, heterozoan carbonates also form in tropical settings, but such heterozoan tropical
sedimentary systems are poorly understood. Nevertheless, distinction between tropical and extratropical heterozoan carbonates
in ancient successions is crucial for accurate paleoenvironmental and paleoclimate reconstructions. Here, surficial Holocene
and Pleistocene sediments of the northern Mauritanian shelf are studied as an example of a tropical eutrophic carbonate
depositional system (11 mg?L21 Chl-a [chlorophyll-a]). Upwelling nutrient-rich waters push onto the wide Mauritanian shelf,
where they can warm up to in excess of 25uC. This condition favors production of heterozoan carbonates dominated by bivalves
and foraminifers, even in this tropical setting. In addition, sediments are provided by eolian input from the desertic hinterland.
The resulting sediments are carbonate and mixed carbonatesiliciclastic facies, in which the carbonates are characterized by a
mixture of tropical and cosmopolitan taxa. Benthic photosynthetic biota are absent while suspension-feeding organisms are
dominant. This foramol grain association on a shelf scale is reminiscent of cool-water carbonates, therefore recognition of
warm-water heterozoan carbonates relies on key taxa related to tropical waters within the biota assemblages associated with a
highly productive environment.

INTRODUCTION

Formation of carbonate sediment is related to biological activity within

the ecosystem of a given depositional environment. Beside temperature,
multiple factors such as salinity, type and availability of substrate,
nutrient concentration, water depth, light penetration, water energy,
seawater chemistry, and siliciclastic supply control the rates and style of
biogenic carbonate production (Hallock and Schlager 1986; Carannante
et al. 1988; Pomar 2001; Mutti and Hallock 2003; Pomar et al. 2004;
Wright and Burgess 2005; Pomar and Hallock 2008; Westphal et al.
2010). In consequence, carbonate sediments are multiparameter archives
of environmental conditions that can be useful for reconstructing
paleoecology and paleoclimate.
In the modern, most tropical settings are characterized by oligotrophic
warm waters. In such settings, carbonates are produced predominantly by
autotrophic biota such as calcareous green algae and mixotrophic biota
(sensu Hallock 1981), such as zooxanthellate corals. Such associations
have been termed chlorozoan by Lees and Buller (1972) and photozoan
by James (1997). In contrast, carbonate sediments from temperate to
polar regions are dominated by heterotrophic biota, corresponding to the
foramol association of Lees and Buller (1972) and the heterozoan
association of James (1997) (e.g., Nelson and Bornhold 1983; Freiwald
* Present Address: Universite de Provence, Laboratoire de Geologie des
Syste`mes et des Reservoirs Carbonates, case 67, 3 place Victor Hugo, 13331
Marseille Cedex 3, France
{ Present Address: Dipartimento di Scienze della Terra, Universita` di Roma
La Sapienza, Ple Aldo 7 Moro, 5. I-00185 Roma, Italy
{ Present Address: Leibniz-Center for Tropical Marine Ecology, Fahrenheitstrasse 6, 28359 Bremen, Germany

Copyright E 2011, SEPM (Society for Sedimentary Geology)

1993; Freiwald and Henrich 1994; Henrich et al. 1995; Rao 1996).
However, foramol or heterozoan associations not only form in cool to
cold-water settings but occur in all climate belts from the poles to the
tropics (Lees and Buller 1972; Lees 1975; Mutti and Hallock 2003; Wilson
and Vecsei 2005). As a consequence, interpreting heterozoan versus
photozoan or foramol versus chlorozoan occurrences as indicative of
cold or temperate versus tropical conditions can result in misleading
paleoclimatic and paleoenvironmental interpretations (cf. Edinger et al.
2002; Pomar et al. 2004).
An increasing number of ancient examples of tropical heterozoan
carbonates have recently been described (Neogene: Brandano and Corda
2002; Pomar et al. 2004; Triassic: Hornung et al. 2007; Pennsylvanian:
Samankassou 2002). Recognition and interpretation of such occurrences
requires detailed facies description, including taxonomic determination of
the skeletal components. In addition to detailed study of the rock record,
modern analog studies provide a means for better understanding of highnutrient tropical settings.
Modern heterozoan carbonate depositional systems in tropical to
subtropical latitudes in most cases are related to oceanographic upwelling
causing elevated nutrient levels (Hallock and Schlager 1986; James 1997;
cf. Westphal et al. 2010). At the same time, the cool upwelling waters
lower the water temperature in tropical seas so that the effect of high
nutrient concentrations is overshadowed by the effect of colder
temperatures and cannot be studied independently (cf. Halfar et al.
2004). In contrast to this general pattern, on the wide shelf off northern
Mauritania cool, upwelling nutrient-rich waters warm to tropical
temperatures, creating a warm-water eutrophic ecosystem. Whereas in
the southern part of this area (the wide Golfe dArguin), carbonate
sediment is diluted by large amounts (up to 63%) of eolian silt, in the
northern part carbonate content reaches up to 93% (Michel et al. 2009).

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J. MICHEL ET AL.

JSR

FIG. 1.Location and bathymetry of the

Golfe dArguin study area between Cap Blanc
and Cap Timiris. Note the large expanse of
shallower water (, 10 mwd) in the center of the
gulf, known as the Banc dArguin. This area
includes water depths of less than 5 m. Largescale surface currents (gray arrows) and dominant wind-induced surface current direction
(large gray arrow) are shown (modified from
Domain 1985; Hanebuth and Lantzsch 2008;
Michel et al. 2009).

In this paper, the facies of this eutrophic tropical sedimentary system are
studied with focus on ecological data to gain a better understanding of
this type of carbonate despositional environment and to improve the
interpretation of ancient counterparts.
STUDY AREA

The narrow (, 65 km) continental shelf of northwest Africa broadens

offshore northern Mauritania to the Golfe dArguin, with a width of
some 150 km (Fig. 1). This extensive gulf hosts the shallow Banc
dArguin, with water depths of less than 10 m and in many areas less than
5 m (Piessens 1979; Sevrin-Reyssac 1993; Wolff et al. 1993). We here refer
to the inner shelf as the area from the coastline to the abrupt 20 m break
in slope, which borders the shallow Banc dArguin. The shelf below this
bathymetric break down to 50 mwd (meters water depth) we refer to as
mid shelf. The mid shelf is about 50 km wide in the northern part of the
study area, whereas it narrows to a few kilometers in width in the south,
where it is cut by submarine canyons. The outer shelf we define here as
the area below 50 mwd down to the shelf break at 100150 mwd.
The waters off northern Mauritania are among the most productive
marine areas in the world (11 mg?L21 Chl-a [chlorophyll-a]; Quack et al.
2007) and are important fishing grounds (Binet et al. 1998). The origin of
the high productivity lies in the elevated nutrient levels. Oceanic upwelling
causes elevated concentrations of phosphate and nitrate on the shelf (up
to 1.3 and 18 mmol.L21 in the surface waters, respectively; Quack et al.
2007), whereas influx of eolian dust is thought to result in elevated iron
concentrations (e.g., Ohde and Siegel 2010).

Oceanic upwelling along the northwest African coastline stretches from

12u N to 33u N (Mittelstaedt 1991; Van Camp et al. 1991), and is reflected
in the presence of cool-water carbonates in most parts of this region
(Summerhayes et al. 1976). Between 20u N and 25u N, oceanic upwelling
occurs year round. Seasonality in the Golfe dArguin, however, induces
an increasing influence of the Guinea Current during summer, related to
latitudinal movements of trade winds that causes sporadic interruptions
of upwelling (Mittelstaedt 1983). Where the upwelling waters enter the
Banc dArguin, minimum temperatures are 16uC. In the shallow Golfe
dArguin, however, the upwelling waters warm to subtropicaltropical
temperatures; on the bank, water temperature exceeds 25uC in summer
and does not drop below 18uC in winter (Sevrin-Reyssac 1993; Quack et
al. 2007). At the same time, the trophic level remains high, resulting in
mesotrophic to eutrophic warm-water conditions.
The Golfe dArguin is characterized by high water energy caused by
swell from the northwest, waves, tides, and wind-driven and deep
currents. Whilst trade winds drive surface currents southwards (speed of
up to 50 cm?s21), undercurrent flows northward along the slope and shelf
break up to 20 cm?s21 (Shaffer 1974; Mittelstaedt 1991). On the
northwest African shelf, tidal currents are usually weak; they are strong
only inshore, as on top of the Banc dArguin (up to 150 cm?s21 in narrow
channels; Piessens 1979; Mittelstaedt 1991). Wind-generated waves are
thought to influence the seafloor on the shallow bank (Piessens 1979).
Swell from north or northwest displays wave lengths of 250 m (measured
on aerial photos) and is thought to play a major role in sediment
remobilization off northwest Africa (cf. Summerhayes et al. 1976;
Piessens 1979).

JSR

TROPICAL HETEROZOAN CARBONATE

643

FIG. 2.Distribution of carbonate contents

of bulk sediment surface samples in the
Golfe dArguin.

The sedimentary system of the Golfe dArguin is characterized by

mobile sand waves in the northern part, including the Baie du Levrier.
This coarse-grained sediment is largely biodetrital, and carbonate
contents reach up to 93% (Michel et al. 2009). On the Banc dArguin,
most material consists of skeletal sand and in coastal parts locally of
carbonate-poor sediments (Piessens and Chabot 1977; Piessens 1979). In
the Baie de Saint-Jean, where hypersaline conditions occur, and in rare
cases in coastal regions of the Baie du Levrier, ooids are present
(Koopmann et al. 1979; Stein 1980). Along the pronounced bathymetric
break of the shallow Banc dArguin, to the outer, deeper part of the Golfe
dArguin, bioclastic sandy material accumulates. Towards the south,
finer-grained sediment dominates that is largely composed of eolian dust,
and carbonate contents are below 50% (Michel et al. 2009). In the
southernmost part of the Golfe dArguin, the shelf is incised by a series of
small canyons, which on the slope merge to the Timiris canyon (Shaffer
1974; Krastel et al. 2004).
MATERIAL AND METHODS

The samples studied here were collected during two cruises of R/V
Poseidon (Westphal et al. 2007; Zonneveld et al. 2010) Sampling of the
shallow Banc dArguin was not feasible during these cruises because of
the shallow water depth (, 10 m) and migrating bedforms that
prohibited the research vessel from entering this area. Sampling therefore
took place on the off-bank shelf, based on the consideration that in a softbottom bioclastic sedimentary system information on the updip

carbonate production is recorded in the downdip sediment as a result

of downslope transport.
A total of 74 surface samples were taken with a van Veen grab and a
box corer (Fig. 2). Van Veen grab recovers materials up to 20 cm below
the sedimentwater interface. Study of the box cores was restricted to the
surface sediment layers. The sediment is strongly bioturbated and thus
considered homogeneous, and no attempt was made to quantify live and
dead shells apart from a visual estimate; the whole resulting assemblages
are studied qualitatively for environmental interpretation. Carbonate
content as a percentage of the bulk sediment is determined using the
Carbometer method, i.e., by measuring the increase in gas pressure after
reacting the ground sample with HCl (Muller and Gastner 1971).
Multiple measurements lead to an internal error of less than 1%. The
carbonate mineralogical composition of 74 samples was determined using
a Bragg-Brentano X-ray diffractometer (XRD). The percentage of the
carbonate minerals (aragonite, high-Mg calcite 5 HMC, minimum of
4 mol. % MgCO3, and low-Mg calcite 5 LMC, maximum of 4 mol. %
MgCO3; see Flugel 2004) relative to total carbonate content was
calculated from peak area ratios using calibration curves. Noncarbonate
components were not further investigated.
For grain-size analyses, bulk samples (n 5 61) were wet sieved at
63 mm. The coarse-grained fraction was split into different sub-fractions
with a sonic sifter (63125 mm, 125250 mm, 250500 mm, 5001,000 mm,
and . 1,000 mm, corresponding to very fine, fine, medium, coarse, and
very coarse-grained sand and gravel, respectively). Further dry sieving
was performed on the bulk material of selected samples to determine the

644

JSR

J. MICHEL ET AL.

respective amounts of the very coarse-grained sand (1,0002,000 mm) and

gravel (. 2,000 mm) fractions.
The sedimentary components of 39 samples were analyzed quantitatively to determine composition. Sorting was determined on thin sections
of bulk sediments. These samples are from the shallow subtidal zone (10
50 mwd) along the Banc dArguin (n 5 34), the top of the bank (n 5 1),
and from deeper water settings (70155 mwd, n 5 4). Samples from the
deeper shelf (. 155 mwd) and the Arguin and Timiris mud wedges were
not analyzed for composition. A minimum of 300 components per grainsize fraction . 125 mm was determined, and included tallies of: red algae,
planktonic foraminifers, benthic foraminifers, bivalves, gastropods,
pteropods, scaphopods, bryozoans, decapods, barnacles, ostracods,
solitary corals, echinoderms, alcyonarian spicules, sponge spicules,
serpulids, aggregated worm tubes, fish remains, fecal pellets, carbonate
clasts (aggregate grains including organically aggregated grains and
intraclasts), unidentified bioclasts (biogenic grains of non-determinable
origin), and siliciclastic grains (mostly quartz). Mollusks and foraminifers
were classified to species level where possible for obtaining a reliable
environmental interpretation of the sedimentary facies. For the determination of the mollusks, the systematics of Nikle`s (1950), Gofas et al.
(1985), Cosel (1995, unpublished data), and Ardovini and Cossignani
(2004) was used; for foraminifers, the systematics of Colom (1950, 1974),
Loeblich and Tappan (1987), Colom and Mateu (2000), and Ellis and
Messina (1940) was employed.
Sedimentary facies were determined on the basis of grain size and
composition of the sediment. Hierarchical cluster analysis (dendrogram
using Wards method and the Euclidean distance measure based on
percentages of grain-size fractions, carbonate content, and composition)
was performed to statistically distinguish groups of samples using PAST
version 2.01 for Windows (Hammer et al. 2001). Non-determinable grains
were not included in the statistical analysis. These unidentified grains,
however, are considered in carbonate content and grain-size data; they
are further taken into account for sedimentary interpretations (i.e.,
hydrodynamics and time averaging).
Radiocarbon ages were determined for mollusk shells that were
strongly bored, physically eroded, and stained, on the basis of the
assumption that they have been reworked and may represent maximum
ages of the material in the surface sediment. The measurements were
undertaken by AMS 14C at the University of Poznan, Poland. The raw
14
C dates were calibrated using CALIB version 5.0.1 (Stuiver and Reimer
1993) and the calibration curve Marine04 incorporating a reservoir-age
correction of 400 years (Hughen et al. 2004).
RESULTS

siliciclastic-rich (for the grain-size distribution in the silt fraction, see

Michel et al. 2009).
Component Analysis of the Loose Sediment
The clear dominance of mollusks and foraminifers in most samples
(Fig. 3) places the sediment of the Golfe dArguin in the foramol grain
association of Lees and Buller (1972) and the heterozoan association of
James (1997). Bivalves, small benthic and planktonic foraminifers,
barnacles, and echinoderms are the most common bioclasts in the
sediments (Fig. 3). Bivalve shells are the most abundant carbonate grains
in 33 out of 39 samples and in none of the samples constitute less than
19% of the identified bioclastic grains (Tab 1; in the following,
abundances of identified bioclasts are given as relative percentages of
bioclastic grains excluding unidentified bioclasts and siliciclastic grains).
Of the six samples with less than 30% bivalves, one from the Baie du
Levrier is dominated by barnacle fragments and five from the southern
part of the Golfe dArguin are dominated by benthic foraminifer tests
(Fig. 3). Small benthic foraminifers are present in most parts of the gulf
and are most abundant in the southern part of the study area. Planktonic
foraminifers constitute . 5% of the identified bioclasts in nine samples,
four of which are from the outer shelf and five are from the mid shelf.
Barnacle fragments are present predominantly in the shallowest samples
in the northern part of the Golfe dArguin and on the Banc dArguin.
Echinoderm fragments are common, but abundance . 5% is restricted to
the Baie du Levrier and the mid shelf.
Less common bioclasts include gastropods, which are ubiquitous but
exceed 5% of the identified grains in only six samples, four from the
southernmost part of the area. Scaphopods are rare. Aggregated wormtube fragments show abundances . 5% in the Baie du Levrier and on the
southern mid shelf. Crustaceans other than barnacles are rare and include
decapods and ostracods. Bryozoan fragments occur in all samples but do
not exceed 5% of the identified bioclasts. Fish remains, fecal pellets,
organically aggregated grains that might represent fragments of
aggregated worm tubes, and sponge spicules are rare, while alcyonarian
spicules, serpulids, pteropods, and ahermatypic coral fragments are very
rare. Fragments of red algae and intraclasts (counted as aggregates in
Table 2) were observed exclusively in the sample from the shallow Banc
dArguin.
Each mollusk and foraminifer assemblage is composed of a mixture of
cosmopolitan and tropical species. Most bioclasts consist of fragmented
shells. Unidentified bioclasts range between 1 and 42% of the sediment
grains . 125 mm (Tab 1). The abundance of siliciclastic grains is highly
variable and ranges from 0 to 70% of the bulk of the identified grains.
Most siliciclastic grains are composed of quartz.

Carbonate Content and Grain Size

The carbonate content in the sediments of the Golfe dArguin ranges
from 35 to 93% (n 5 74; 53 samples . 50%; Fig. 2, Tab 1). The
sedimentary system thus is a mixed carbonatesiliciclastic system; it
includes a carbonate-dominated area in the northern study area, where
carbonate content reaches values up to 7093%. In the vicinity of Cap
Blanc, carbonate content locally is lower (4462%) as a result of quartz
sand dunes migrating into the sea. With increasing water depth,
carbonate contents decrease from . 70% to , 50% before increasing
again between 100 and 200 mwd to . 80%. The central study area is
characterized by intermediate values between 4570%, whereas in the
southern part values of 3550% are typical.
Grain-size distribution shows a pattern similar to that of the carbonate
content (Fig. 3, Tab 1). In the north, the most abundant grain sizes are
medium- to coarse-grained sand and even coarse-grained sand to gravel
south of Cap Blanc. In the southern part of the study area, mud and very
fine- to fine-grained sand dominate. The coarse-grained fraction is
composed of biogenic carbonate components whereas the silt fraction is

Mineralogical Analysis of the Carbonate Fraction

Aragonite constitutes 2181% of the bulk sediment (mean of 42%) and
4594% of the total carbonate portion (mean of 68%) and thus is the
dominant carbonate mineral (Fig. 4). Only four samples contain less than
50% aragonite in the total carbonate portion. The highest aragonite
content is found (1) in shallow water depths along the Banc dArguin and
(2) at greater water depths around the shelf break. The lowest aragonite
contents in the carbonate fraction are found in the vicinity of Cap Blanc
between 15 and 40 mwd, in an area located between 35 and 65 mwd off
the northern Banc dArguin, and in four isolated samples farther south
(Fig. 4).
LMC averages at 13% (335%) of the bulk sediment and 23% (551%)
of the total carbonate. Only three samples located directly south of Cap
Blanc contain more LMC than aragonite. HMC constitutes an average of
5% (021%) of the bulk sediment and 8% (041%) of the total carbonate.
The highest HMC content is present in a sample from the central open
shelf.

RAL

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GeoB

11511B
11511C
11511D
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11526
11527
11528
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11549
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11595
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11614

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4.8
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32.5
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25.7
44.7
25.2
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1.9
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4.4
7.8
17.0
9.3
13.3
0.8
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5.8
8.6
1.4
4.0
4.3
12.1
35.0
14.1
25.6
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37.5
28.4
33.9
3.8
0.0
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6.7
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10.3
14.1
12.2
20.2

PFOR BFOR

40.3
46.3
32.7
29.1
51.4
47.7
63.1
46.3
71.7
40.8
65.9
47.3
67.6
39.6
19.3
58.2
61.8
69.7
64.6
75.3
59.4
89.7
45.1
19.2
64.1
67.6
34.4
19.5
19.2
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12.7
83.2
78.2
22.3
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11.5
47.7
50.4
45.5

BIV
1.5
5.9
3.5
0.1
2.3
3.4
2.5
1.5
2.7
3.1
2.3
5.4
1.6
1.3
0.6
3.0
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2.9
1.2
4.3
4.0
2.8
1.7
2.2
0.8
1.7
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6.2
5.5
9.9
1.7
1.4
1.8
1.7
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3.2

2.4
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15.1
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0.7

BAR
3.6
1.0
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2.1
4.2
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OST
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0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.9
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

COR
4.9
6.6
6.1
0.3
0.3
5.5
13.5
0.3
1.0
0.2
0.6
13.3
4.9
15.2
2.5
1.0
0.2
11.6
16.7
2.7
8.0
0.8
2.2
0.0
3.3
1.2
1.0
9.2
18.4
2.9
1.0
0.9
0.9
1.6
5.7
5.7
2.2
2.8
1.5

ECH
4.8
0.2
0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.6

ALC
1.2
0.4
4.9
0.0
0.0
1.3
0.0
0.0
0.0
0.0
0.6
0.0
0.0
0.4
0.0
0.0
0.0
0.4
0.1
0.0
0.1
0.0
0.3
0.0
0.3
0.0
0.0
2.9
0.7
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

SPO
0.1
0.0
0.0
0.0
1.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.3
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.1
0.0
1.0
0.0
0.0
0.4
0.0
0.0
0.0
0.0
0.4
0.0
0.0
0.0
0.0
1.8
0.0
0.6

SER
0.0
5.7
12.5
1.9
0.0
5.8
3.6
0.2
0.2
0.9
0.0
8.3
0.0
2.2
0.0
0.1
0.4
4.2
3.0
0.7
1.3
0.0
0.6
0.1
0.1
0.3
0.0
5.2
5.7
2.8
0.1
0.0
0.1
1.2
4.7
0.1
0.0
0.1
1.8

WRM
0.1
0.0
0.0
0.1
1.2
0.0
0.2
0.0
0.0
0.0
0.0
0.1
0.4
0.0
0.0
0.0
0.0
0.0
0.5
0.2
1.8
0.3
0.1
8.8
1.1
0.1
0.0
0.2
0.1
0.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
3.7
0.7

FSH
0.4
0.8
2.3
4.9
1.6
1.1
0.2
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.5
0.0
0.0
0.0
0.0
0.8
0.6
0.0
0.0
0.0
0.2
0.0
3.1
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

0.4
0.0
0.0
0.0
0.0
0.0
0.2
0.3
0.1
0.0
2.4
1.1
0.0
0.0
0.8
0.5
0.4
0.0
0.2
0.5
1.7
0.0
0.2
0.5
0.1
0.7
8.7
0.0
0.0
0.4
0.0
0.1
0.1
0.3
0.3
0.3
0.9
0.1
2.1

3.6
14.5
5.2
6.9
5.4
11.7
2.5
44.1
7.4
21.5
8.7
8.4
2.2
30.3
60.1
14.5
21.1
1.8
2.6
6.4
10.3
0.4
2.7
1.5
1.1
0.7
29.8
12.1
11.5
0.0
70.1
2.6
6.7
52.9
33.2
61.5
19.4
3.1
7.7

36.5
44
56
56
56
62.5
84
56
83
74
81
77
82
44
55.5
78
71
71
85
87
82
93
69
53
91
81
50
39
39
50
55
75
85
62
45
51
51
85
62.5

0.0
0.1
0.0
1.3
0.0
0.6
5.7
4.4
22.1
5.5
0.6
13.9
1.3
1.5
1.2
23.2
18.0
1.5
2.6
9.5
4.2
4.1
0.2
0.2
0.4
0.7
5.9
0.1
0.1
5.1
0.9
13.4
21.3
3.2
1.0
0.3
1.9
0.6
2.3

92.1
88.5
61.1
66.7
88.8
94.0
89.5
95.1
77.0
94.3
90.9
84.7
97.2
95.5
98.7
75.7
81.7
78.4
92.9
90.0
93.9
95.7
87.2
78.5
98.0
98.5
94.1
30.9
28.8
69.4
98.9
86.2
78.5
96.0
55.6
99.3
88.2
98.0
88.3

7.9
11.5
38.9
31.9
11.2
5.4
4.8
0.5
1.0
0.2
8.5
1.4
1.5
3.0
0.1
1.1
0.3
20.1
4.5
0.5
1.9
0.3
12.7
21.3
1.6
0.8
0.0
69.0
71.1
25.6
0.1
0.4
0.2
0.7
43.3
0.5
9.9
1.4
9.3

FEC AGG QUA Carb Grav Sand Mud

TABLE 1. Data from the 39 component-analyzed samples from the Golfe dArguin used for the cluster analysis showing loose-sediment quantification as percentages (excluding unidentified
bioclasts), carbonate content (dry wt%), and grain-size fractions (gravel, sand, and mud; dry wt%). RAL, red algae; PFOR, planktonic foraminifers; BFOR, benthic foraminifers; BIV, bivalves;
GAST, gastropods, PTER, pteropods; SCAP, scaphopods; BRY, bryozoans; DEC, decapods; BAR, barnacles; OST, ostracods; COR, ahermatypic corals; ECH, echinoderms; ALC, alcyonarian
spicules; SPO, sponge spicules; SER, serpulids; WRM, aggregated worm tubes; FSH, fish remains; FEC, fecal pellets; AGG, aggregates; QUA, quartz grains.

JSR
TROPICAL HETEROZOAN CARBONATE
645

646

JSR

J. MICHEL ET AL.

FIG. 3.Distribution of sediment grain sizes

(bulk) and components (grains . 125 mm) of
surface samples in the Golfe dArguin. BIV,
bivalves; RAL, red algae; PFOR, planktonic
foraminifers; BFOR, benthic foraminifers; BAR,
barnacles; ECH, echinoderms; Other, other
bioclasts; QUA, quartz grains.
14

SEDIMENTARY FACIES

C Ages

The mollusk shells considered to be reworked relict material show

increasing ages with increasing water depth (Tab 2). In less than 40 mwd,
ages reach a maximum of 300 cal yr BP, whereas in 50100 mwd ages
between 9,500 and 15,500 cal yr BP occur.

Observation of the seafloor sediments reveals five sedimentary facies

(Tab 3). This facies definition is defined by statistical analysis based on
grain size, carbonate content, and grain association (Fig. 5). Whereas the
distribution of two facies (F1 and F3) is restricted to shallow water

TABLE 2.Radiocarbon measurements and age calibration using Marine04 curve and a 400-years reservoir age of highly abraded mollusk shells from
surface samples (mwd 5 meters water depth; Hughen et al. 2004).

Lab No.

Sample

Sample depth (mwd)

Material

Poz-31069
Poz-31070
Poz-31073
Poz-26879
Poz-26880
Poz-26881
Poz-26882

GeoB11501
GeoB11501
GeoB11513
GeoB11613
GeoB11614
GeoB13018
GeoB13019

16.5
16.5
35.5
103.3
72.3
37.0
53.0

Donax burnupi valve

Bivalve piece
Gastropod piece
Bivalve piece
Bivalve piece
Bivalve piece
Bivalve piece

14

C ages [14C yr BP]

Modern
525 6 30
600 6 30
13350 6 70
8870 6 50
640 6 30
10250 6 50

1s calibrated
[cal yr BP]

Intercept [cal yr BP]

Modern
118238
149290
1512715426
94669565
255310
1117911262

Modern
180 6 60
220 6 70
15275 6 150
9515 6 50
285 6 30
11220 6 40

JSR

TROPICAL HETEROZOAN CARBONATE

647

FIG. 4.Distribution of aragonite contents in

the Golfe dArguin as percentages of total
carbonate. Aragonite, LMC (, 4 mol.%
MgCO3), and HMC (. 4 mol.% MgCO3)
percentages of the 74 samples are displayed in
the ternary plot showing the overwhelming
dominance of aragonite.

TABLE 3. Sedimentary facies (F) of the Golfe dArguin (Mauritania) based on a cluster analysis including grain size (dry wt%), carbonate content
(CaCO3; dry wt%), and sediment composition of the samples.

FACIES

SAMPLE

TEXTURE (%) AND

SORTING

GeoB11515, GeoB11516, GeoB11528,

GeoB11529, GeoB11532, GeoB11534,
GeoB11601, GeoB11602
GeoB11513, GeoB11522, GeoB11524,
GeoB11525, GeoB11530, GeoB11531,
GeoB11533, GeoB11547, GeoB11549,
GeoB11613
GeoB11514, GeoB11526, GeoB11527,
GeoB11591, GeoB11597, GeoB11603,
GeoB11606

gravel: 06; sand: 9499;

mud: 03
moderately to well sorted

F4
Dust-influenced

GeoB11511B, GeoB11511C, GeoB11511F,

GeoB11511G, GeoB11535, GeoB11607,
GeoB11614

gravel: 02; sand: 8794;

mud: 513
very poorly to well sorted

F5
Mud-rich

GeoB11511D, GeoB11511E, GeoB11540,

GeoB11593, GeoB11594, GeoB11595,
GeoB11604

gravel: 05; sand: 2979;

mud: 2171
poorly to moderately sorted

F1
Donax burnupi
F2
Bivalve

F3
Dune-influenced

gravel: 423; sand: 7696;

mud: 01
moderately to well sorted
gravel: 014; sand: 7899;
mud: 020
poorly to moderately sorted

SEDIMENT COMPOSITION
Quartz and barnacle-rich, bivalve-dominated
Variably fragmented and abraded shells
Clean carbonate sand and gravel
Foraminifer-rich, bivalve-dominated, with variable
amount of echinoderms
Highly fragmented, abraded, and bioeroded shells
Clean carbonate sand
Barnacle-rich, quartz- and bivalve-dominated, with
variable amount of benthic foraminifers and
echinoderms
Variably fragmented and abraded shells
Clean mixed carbonate-siliciclastic sand
Quartz-rich, benthic foraminifer- and bivalve-dominated,
with variable amount of planktonic foraminifers
Mixed abraded-fresh shells
Mixed carbonate-siliciclastic muddy sand to sand
Bivalve- and foraminifer-dominated, with variable
amount of echinoderms and quartz
Mixed abraded-fresh shells
Mixed carbonate-siliciclastic sandy mud to muddy sand

CaCO3
(%)
7193
7191

4462

3769

3956

648

J. MICHEL ET AL.

JSR

FIG. 5.Dendrogram of hierarchical cluster

analysis based on carbonate content, grain size
(mud, sand, and gravel; dry wt%), and composition (excluding unidentified bioclasts) from the
loose-sediment analysis that statistically groups
similar samples. The determination of the
threshold, which defines five groups of samples
and thus the five facies (F15), is based on
subjective observation of bulk sediment.

depths, three facies (F2, F4, and F5) include deposits from a range of
water depths (i.e., mid and outer shelf; Fig. 6). The facies distribution
thus is not predominantly bathymetrycally defined but rather represents a
facies mosaic (i.e., patches on the shelf). An environmental interpretation
of the facies is based on the detailed taxonomical and ecological analysis
of mollusks and foraminifers.

Gari jousseaumeana, both of which have a clear tropical affinity (cf. Cosel
1995). The low number of living organisms is interpreted to indicate that
carbonate production occurs updip of the sampling locations (i.e., on the
outer part of the Banc dArguin), from which the shells are shed and
reworked. Quartz grains are supplied by migrating sand dunes of onshore
Cap Blanc.

Coarse-Grained Donax burnupi-Dominated Sand Facies (F1; Fig. 7A)

Bivalve Fragment-Dominated Sand Facies (F2; Fig. 7B, C)

Carbonate content of these sediments ranges from 71 to 93% (Fig. 6,

Tab 3). Bivalves dominate (especially Donax burnupi), along with
barnacle fragments. These constituents are reflected in the coarse grain
sizes, reaching from medium-grained sand to gravel; sorting is moderate
to good. The carbonate grain association corresponds to a bimol
skeletal assemblage sensu Hayton et al. (1995). Quartz grains are variably
abundant (019% of the grains . 125 mm). This facies is present in
sediment in the shallow northern part of the Golfe dArguin (1723 mwd)
and in two samples located farther south at around 35 mwd (Fig. 6).

The sediments consist of poorly to moderately sorted carbonate

medium and coarse-grained sand (7191% carbonate content) (Fig. 6,
Tab 3). The grains of this facies are highly fragmented, abraded, and
bioeroded (Fig. 7B, C). Besides bivalve fragments, benthic foraminifers
are common. Planktonic foraminifers are common in three samples
(Tab 1), two on the outer shelf. Echinoderm fragments range from 1 to
17% of the identified bioclasts. This facies is a bimol skeletal
assemblage sensu Hayton et al. (1995). It is present on the mid shelf
and on the outer shelf in the northern part of the Golfe dArguin (Fig. 6).

Interpretation and Environmental Conditions.The environment of this

Donax burnupi-dominated facies closely follows the ecological requirements of the genus Donax, which prospers in high-energy, subtidal
environments (cf. Ansell 1983). In the Golfe dArguin, these conditions
are associated with the swell from the northwest. The high trophic
resources (i.e., phytoplankton) related to upwelling, and tropical to
subtropical climate, further provide preferable conditions for this bivalve
(cf. Ansell 1983). In addition to the dominant D. burnupi shells, the
mollusk assemblage includes the bivalve species Crassatina marchadi and

Interpretation and Environmental Conditions.This bivalve fragmentdominated sand facies is present in two different settings, on the mid shelf
and on the outer shelf. The main difference between the mid-shelf
(Fig. 7B) and outer-shelf (Fig. 7C) deposits relate to the grade of
abrasion of the grains, which is more intense on the outer shelf, and to the
taxa present. In the mid-shelf deposits, Donax burnupi fragments
dominate. In the outer-shelf sediments, shallow-water-related Ervilia
castanea fragments (cf. Morton 1990) dominate, whereas the betterpreserved mollusk shells are of deep-water origin (e.g., Anodontia

JSR

TROPICAL HETEROZOAN CARBONATE

1615

20 m

50 m

17W

100 m

21N

1000 m

1745

649

Cap
Blanc

foramol-type
sediments
Seagrass
20N

Ooids
)
Cap
Timiris

Component-analyzed sample

1915

F1
Coarse
sand

F2
Bivalve
fragment
sand

F3
Quartz
bivalve
sand

F5
F4
Siliciclastic Siliciclastic
biv-foram biv-foram
muddy
sand

Components
QUA BIV
Other

RAL

ECH

PFOR

BAR

500-1000
125-250
<63
70-95

70-90

45-60

35-70

BFOR
>1000
250-500
63-125

40-55

FIG. 6.Facies distribution in the study area. Interpolation between locations of samples on which detailed component analysis was performed is based on facies
analysis of other samples and on previous sedimentological data from Domain (1985). Average component composition (. 125 mm fraction), grain sizes, and carbonate
content range of facies are given, as well as literature data for inner-shelf sediment.

senegalensis, Mesalia flammifera). The 14C date of highly abraded bivalve

material at 103 mwd indicates an age of 15.5 cal kyr BP (Tab 2). Thus,
these outer-shelf deposits are interpreted as palimpsest sediments, where
relict grains have remained exposed on the seafloor for thousands of
years.
The mollusk assemblage of this bivalve-fragment-dominated sand
facies is a mixture of cosmopolitan and northwest African taxa. The
endemic malacofauna includes both bivalves and gastropods (e.g., the
northwest African species Mesalia mesal and Turritella bicingulata; cf.

Ardovini and Cossignani 2004); some clearly point to a tropical

environment (e.g., Modiolus nicklesi, Venus erronea, Persicula cingulata,
and Persicula cornea; cf. Nickle`s 1950).
The foraminiferal assemblage is composed of a mixture of small
benthic and planktonic taxa. The benthic association is dominated by
sediment dwellers such as Pararotalia sp., non-keeled Elphidium spp.,
miliolids, and textulariids. In addition, species such as Cibicides refulgens,
Elphidium macellum, Lobatula lobatula, and Miniacina miniacea are
abundant, generally associated with phytal substrates (cf. Langer 1993).

650

J. MICHEL ET AL.

JSR

JSR

TROPICAL HETEROZOAN CARBONATE

Rare taxa (i.e., Cassidulina laevigata, bolivinids, buliminids, and

uvigerinids) are related to low-oxygen conditions (cf. Bernhard and Sen
Gupta 1999). Most of the benthic taxa present are ubiquitous and thus
are not indicative of particular environmental conditions; however, the
genus Pararotalia indicates warm climatic conditions (cf. Murray 2006).
The planktonic taxa are mostly deep-water species (sensu Be 1977) that
live at depths greater than 100 m (Globorotalia inflata and Neogloboquadrina pachyderma). However, the epipelagic (, 50 mwd, sensu Be 1977)
Globigerinoides trilobus is also abundant. Regarding their climatic
significance, the planktonic foraminiferal association consists of an
assemblage of cold (e.g., Globigerina bulloides and N. pachyderma), but
mostly transitional (e.g., G. inflata) and subtropical to tropical (i.e.,
Globigerina calida, Globorotalia crassula, Globorotalia menardii, and
Pulleniatina obliquiloculata) taxa. This group is interpreted to reflect the
complex oceanographic situation of the region and is in agreement with
previous observations of the living planktonic foraminiferal assemblage
(Miro 1973; Thiede 1975a, 1975b; Mateu 1979).
Quartz-Rich Bivalve Sand Facies (F3; Fig. 7D, E)
Sediments are moderately to well sorted and are dominated by fine- to
coarse-grained sand-size grains. Quartz is abundant and dilutes the
carbonate content, which ranges from 44 to 62% (Fig. 6, Tab 3). The
most common bioclasts are bivalves, along with barnacles. Some whole
barnacle shells are still attached to their substrate, such as bivalve shells.
Small benthic foraminifers are more abundant in the samples dominated
by fine- to medium-grained sand from the close vicinity of Cap Blanc
(Fig. 3). Echinoderm fragments range from 0 to 22% of the identified
bioclasts. This mixed carbonateterrigenous material is variably fragmented and abraded and is dominated by Donax burnupi shells close to
Cap Blanc (Fig. 4B) and by several other bivalve species on top of the
Banc dArguin (e.g., Veneridae; Fig. 4C). Red algae fragments and
intraclasts are present on top of the Banc dArguin.
Interpretation and Environmental Conditions.The environment of this
mixed bivalve-quartz facies is similar to that of the Donax burnupi facies
(i.e., shallow subtidal and swell-exposed), but this facies is additionally
influenced by quartz sediment from Cap Blanc. Also, this quartz-rich
facies is dominated by finer grain sizes (i.e., fine- to medium-grained
sand). The accumulation of quartz probably corresponds to a submerged
extension of Cap Blanc, as evident from the seafloor morphology (Fig. 1).
On top of the Banc dArguin in the southern part of the Golfe dArguin,
quartz grains are also abundant but coarser (i.e., medium- to very coarsegrained sand). The very shallow water of the Banc dArguin (4 mwd) is
the only location in the Golfe dArguin where photic-related benthic
carbonate grains (i.e., red algae fragments) were found (see also Piessens
1979).
Suspension-feeding organisms (e.g., barnacles and the bivalve species
D. burnupi) overwhelmingly dominate the bioclast composition. Warmwater-related taxa include the tropical bivalve Pitar belcheri and the
tropicalsubtropical gastropods Marginella senegalensis and Persicula
blanda in the vicinity of Cap Blanc, and the bivalves Carditamera contigua
and Diplodonta dautzenbergii on the Banc dArguin. The shallow

651

subtidal-related bivalves Dosinia exoleta and Carditamera contigua are

present. Benthic foraminifers include sediment dwellers (e.g., non-keeled
Elphidium spp. and miliolids) and species associated with phytal
substrates (sensu Langer 1993) such as Cibicides refulgens and Elphidium
crispum.
Fine-Grained SiliciclasticBivalveForaminifer Sand Facies (F4;
Fig. 7F, G)
The dominant grain size of this facies is very fine- and fine-grained sand
with minor amounts of mud and extremely rare gravel (Fig. 6, Tab 3).
Sediments are poorly to well sorted; the sample from the Baie du Levrier
is poorly sorted. Carbonate content ranges from 37% to 69%. The
carbonate grain association corresponds to foramol sensu stricto:
bioclasts consist mostly of mollusks (bivalves) and foraminifers. The
content of quartz grains ranges from 2 to 14% of the fraction . 125 mm.
This facies is found on the mid shelf in the southern part of the Golfe
dArguin, in the Baie du Levrier, and on the northern outer shelf (Fig. 6).
Interpretation and Environmental Conditions.As for the bivalvefragment-dominated sand facies, the fine-grained siliciclasticbivalve
foraminifer sand facies is found in various settings of the shelf. This
mosaic-type distribution of facies is related to two parameters. In situ
carbonate production provides variable grain associations and grain-size
spectra in the Golfe dArguin. Reworking of these clastic carbonates and
of silicilcastics leads to resorting of the sediment according to hydraulic
properties of the grains. For example, fine-grained sediment is deposited
not only below 50 mwd, where wave action is reduced (Fig. 7G) but also
in protected settings of the Baie du Levrier (Koopmann et al. 1979).
The mollusk assemblage consists of a mixture of endemic northwest
African taxa and cosmopolitan species. Some bivalve species (e.g.,
Cardiocardita ajar and Tellina densestriata) and gastropod species (e.g.,
Periscula cingulata and Prunum annulatum) show a clear tropical affinity
(cf. Nickle`s 1950; Cosel 1995). Other taxa, especially bivalves such as
Cuna gambiensis and Donax burnupi are well adapted to high-nutrient
environments such as upwelling areas (cf. LeLoeuff and Cosel 1998;
Branch et al. 2002).
The foraminiferal assemblage consists mostly of benthic species that
correspond to sediment dwellers (e.g., Cancris auriculus, non-keeled
Elphidium spp., miliolids, and textulariids) and species associated with
phytal substrates (e.g., Cibicides refulgens, Lobatula lobatula, Miniacina
miniacea, Planorbulina mediterranensis, and Rosalina macropora; cf.
Langer 1993; Murray 2006). Some other benthic taxa are related to
low-oxygen conditions (e.g., Bolivina sp. and Textularia spp.; cf.
Bernhard and Sen Gupta 1999) or deep-water environments (i.e.,
Planulina ornata; cf. Colom 1974).
In contrast, planktonic species are much scarcer and are dominated by
epipelagic and mesopelagic taxa (e.g., Globigerina spp. and Globigerinoides spp.). As for the other facies, cold-water species (e.g., Globigerina
bulloides) coexist with transitional (e.g., Globorotalia inflata) and tropical
to subtropical (i.e., Globorotalia crassula and Globorotalia menardii) taxa.
This situation is interpreted to reflect upwelling of deep, cool water
masses, mixing at the surface with warm water masses.

r
FIG. 7.Representative bulk sediments of the study area: A) F1, coarse-grained Donax burnupi-dominated sand facies, carbonate sand and gravel, GeoB11515,
21 mwd; B) F2, bivalve fragment-dominated sand facies, carbonate sand, GeoB11547, 30 mwd; C) F2, bivalve fragment-dominated sand facies, relict carbonate sand,
GeoB11613, 103 mwd; D) F3, quartz-rich bivalve sand facies, mixed carbonatesiliciclastic sand, GeoB11603, 25 mwd; E) F3, quartz-rich bivalve sand facies, mixed
carbonatesiliciclastic coarse sand, GeoB11591, 4 mwd; F) F4, fine-grained siliciclasticbivalveforaminifer sand facies, mixed carbonatesiliciclastic muddy sand,
GeoB11511C, 31 mwd; G) F4, fine-grained siliciclasticbivalveforaminifer sand facies, mixed relict carbonatesiliciclastic muddy sand, GeoB11614, 72 mwd; H) F5,
siliciclasticbivalveforaminifer muddy facies, mixed carbonatesiliciclastic sandy mud, GeoB11595, 41 mwd. B, barnacle; Bi, bivalve; Br, bryozoan; D, Donax burnupi; E,
echinoderm; bF, benthic foraminifer; pF, planktonic foraminifer; G, gastropod; Q, quartz; W, aggregated worm tube. Scale is 5 mm for each picture.

652

J. MICHEL ET AL.
SiliciclasticBivalveForaminifer Muddy Facies (F5; Fig. 7H)

Carbonate content of this facies is lowest and ranges from 39 to 56%

(Fig. 6, Tab 3). This facies is characterized by high mud content, very
poor to moderate sorting, and a bivalve- and foraminifer-dominated
biota with the exception of one sample from the Baie du Levrier that is
dominated by barnacle fragments. Subordinate components of the
. 125 mm fraction include echinoderms, aggregated worm tubes, and
quartz. The mud-rich facies is present in the southernmost part of the
study area, on the mid to outer shelf, and in the Baie du Levrier (Fig. 6).
Interpretation and Environmental Conditions.The mollusk assemblage
consists of a mixture of cosmopolitan, endemic northwest African, and
tropical taxa. Bivalves Cuna gambiensis and Tellina densestriata and
gastropods of the family Marginellidae such as Marginella glabella reflect
tropical water temperatures (cf. Nikle`s 1950; Cosel 1995). Mollusks
reflect the muddy substrate (e.g., Nuculana bicuspidata and Tellina
compressa). The occurrence of the bivalves Anodontia sp., Myrtea
spinifera, and Thyasira flexuosa is interpreted to be related to low-oxygen
conditions, which suggest high organic-matter concentrations in the finegrained sediments.
The foraminiferal assemblage includes small benthic and planktonic
taxa. Among the small benthic species, several low-oxygen-tolerant taxa
are present, many of them associated with muddy bottoms (e.g.,
Cassidulina laevigata, bolivinids, buliminids, and uvigerinids). Other
soft-bottom dwellers found in this facies are miliolids, textulariids, nonkeeled elphidiids, and nonionids. The presence of Cibicididae (e.g.,
Cibicides refulgens and Lobatula lobatula) and encrusting forms (i.e.,
Miniacina miniacea and Planorbulina mediterranensis) often reported as
epiphytic forms (cf. Langer 1993) may be related to the occurrence of
phytal substrates (i.e., seagrass) that have been described from inner parts
of the Banc dArguin (Wolff and Smit 1990; Hemminga and Nieuwenhuize 1991).
The planktonic foraminifers are dominated by meso-epipelagic species.
As in other facies, the planktonic foraminiferal assemblage consists of a
mixture reflecting different climatic requirements. Thus, subarctic to
transitional species (e.g., Globigerina bulloides, Globorotalia inflata, and
Neogloboquadrina pachyderma) co-occur with species adapted to subtropical to tropical conditions (e.g., Globigerina calida, Globorotalia
crassula, Globorotalia menardii, Globorotalia tumida, and Pulleniatina
obliquiloculata).

JSR

(Fig. 1) that results in winnowing, net sediment transport to the south,

and deposition of the eolian silt in the southern part of the Golfe
dArguin. This sedimentation pattern is reflected by the overall north
south trend from coarse-grained, carbonate-dominated sediments in the
north to fine-grained, siliciclastic-dominated deposits in the south (Michel
et al. 2009).
The carbonate deposits largely lack a photic-related zonation, because
most of the benthic carbonate-secreting organisms are aphotic. The
bioclastic composition of the five facies therefore is similar throughout,
dominated by shells and fragments of bivalves and foraminiferal tests. As
for purely siliciclastic sediments and overshadowing the influence of
biogenic production on grain sizes, the hydraulic regime largely controls
the grain-size spectra and sorting of the sediment. Thus, the facies
arrangement reflects the interaction between: (1) carbonate production,
controlled by the biology and ecology of the carbonate-secreting biota
(e.g., the large portion of centimeter-size Donax burnupi shells present in
the northern shallow part of the Golfe dArguin; F1, F2, and F3), and (2)
hydrodynamics and topography, which influence dispersion and distribution of the carbonate and noncarbonate sediments (e.g., greater
contents of mud and fine-grained-sand in the southern part of the Golfe
dArguin; F4 and F5; Fig. 6).
Reworking of sediment not only influences facies distribution, the
resulting abrasion and fragmentation partly explains the large proportion
of undeterminable bioclasts (up to 40%). On the outer shelf, most skeletal
coarse grains are relict material that may represent shallow-water
(, 50 mwd), bivalve-rich deposits formed during the last transgression.
The typically low sedimentation rate of open-shelf heterozoan carbonates
(cf. Nelson 1988; James 1997) and the strong current regime (cf. Bein and
Futterer 1977) might have been responsible for the exposure of these
sediments on the seafloor for thousands of years.
The depositional system of the Golfe dArguin became established only
in the Holocene after submergence of the Banc dArguin around 7 kyr BP
(Hanebuth and Lantzsch 2008). The sediment of this depositional system
therefore composes a thin veneer. Seismic data from the shelf off the Banc
dArguin confirm that the Holocene sediment forms a layer of up to some
10 m overlying an erosional surface at the top of the Pleistocene
succession. The abraded relict grains that are dated between 15.5 cal kyr
BP at 103 mwd to modern at 16.5 mwd (Tab 2) reflect the succeeding
postglacial sea-level rise.
IMPLICATIONS: HETEROZOAN WARM-WATER CARBONATES

MINERALOGY OF HETEROZOAN CARBONATES

The anticipation of calcite being the dominant calcium carbonate

mineral in heterozoan carbonates (whereas aragonite dominates photozoan associations, cf. Nelson 1988; James 1997; Flugel 2004) is not
confirmed for the infaunal bivalve-dominated grain association studied
here, which is clearly aragonite-dominated (Fig. 4). Other aragonite-rich
heterozoan associations are known from modern cool-water deposits
from New Zealand (Nelson et al. 1982; Gillespie and Nelson 1997) and
the South Australian Shelf (James et al. 2005). The dominance of
aragonite, together with the high organic content of the sediment, reduces
the preservation potential of the biotic association studied here
considerably (cf. Smith and Nelson 2003; Wright and Cherns 2008).
FACIES DISTRIBUTION AND DEPOSITIONAL SYSTEM

The sedimentary facies in the Golfe dArguin (Fig. 6) form a facies

mosaic rather than bathymetrically defined depositional belts. Eolian
sediment input from the Sahara and Sahel spreads uniformly over the
entire Golfe dArguin (cf. Stuut et al. 2005). The orientation of the Banc
dArguin relative to the swell from the northwest, coupled to the
prevailing winds, leads to a northsouth-directed hydraulic regime

The facies in the study area are dominated by bivalve fragments and
foraminifers and thus are heterozoan carbonates sensu James (1997) and
foramol carbonates sensu Lees and Buller (1972). Whereas this carbonate
depositional system displays facies also typical of nontropical systems
(Tab 4), the oceanographic situation is clearly a warm-water eutrophic
setting. Seawater temperatures on the Banc dArguin (1829uC) are
tropical, and those of the mid and outer shelf (1625uC) correspond to the
boundary of tropical and warm-temperate conditions (cf. Flugel 2004;
Tab 4). The carbonate mineralogy is aragonite-dominated, and ooids are
found in isolated coastal environments (e.g., Baie de Saint-Jean; Stein
1980). The low-latitude conditions on the open shelf of the Golfe
dArguin are indicated throughout the entire shelf by tropical-related taxa
(mollusks and foraminifers) that allow the biota to be distinguished from
temperate counterparts.
Thus, the attributes of the Mauritanian shelf sedimentation do not
simply fit into common carbonate classifications, and demonstrate the
multidimensional ecological control of carbonate sedimentation. The
Golfe dArguin thus might serve as modern analog for ancient carbonate
sedimentary systems such as Paleozoic upwelling ramps (James 1997;
Martindale and Boreen 1997), Cretaceous warm, high-nutrient molluskdominated systems (e.g., Carannante et al. 1995; Allmon 2007) or

JSR

TROPICAL HETEROZOAN CARBONATE

653

TABLE 4. Comparison of environmental and facies attributes characterizing tropical, subtropical, temperate-polar, and eutrophic tropicalsubtropical
(Mauritanian-type) carbonates based on Nelson (1988), James (1997), and results of the present study.
Environmental and facies
parameters

Subtropical carbonates

Carbonate association
Latitude
Depositional setting
Terrigenous supply
Mean water temperature
Trophic conditions
Photic conditions (shallow
water)
Reefs
Carbonate content

photozoan
between 30 u N and 30 u S
rimmed shelves or platforms
low
. 22 uC
oligotrophic
euphotic

photozoan/ heterozoan
between 30 u N and 30 u S
open shelves or ramps
low
1822 uC
oligotrophic to mesotrophic
oligophotic to euphotic

heterozoan
beyond 30 u N and 30 u S
open shelves or ramps
low to high
, 18 uC
mesotrophic to eutrophic
oligophotic to aphotic

heterozoan
between 30 u N and 30 u S
open shelves or ramps
low to high
1822 uC (1629 uC)
eutrophic
aphotic to oligophotic

abundant
very high (. 90%)

rare
very high (. 90%)

absent
moderate to very high (50100%)

Ooids
Major skeletal carbonate
components

common
hermatypic corals, benthic
foraminifers, mollusks,
calcareous green and red
algae

rare or absent
calcareous red algae, large
benthic foraminifers,
bryozoans; few hermatypic
corals

Terrigenous material
Shell preservation
Carbonate mineralogy

rare
generally good
aragonite dominant

Diagenetic regime

constructive (grain
preservation and chemical
precipitation)

rare
poor to good
low- and/or high-Mg calcite
dominant
destructive (grain dissolution
and maceration,
biodegradation)

rare or absent
moderate to very high
(50100%)
absent
bryozoans, bivalves, small
benthic and planktonic
foraminifers, barnacles,
echinoderms, serpulids,
brachiopods, sponges,
calcareous red algae
rare to abundant
poor to good
low- and/or high-Mg calcite
dominant
destructive (grain dissolution
and maceration,
biodegradation)

Neogene carbonate ramps formed in nutrient-rich tropical waters (e.g.,

Pomar et al. 2004; Brandano and Corda 2002). The depositional
environment of the Golfe dArguin is a tropical broad shelf influenced
by upwelling. The nutrient-rich waters lead to increased plankton
productivity and subsequently to high turbidity, which is further
increased by dust input from the Sahara (Fig. 8). In addition, the
seasonal variability following the southern boundary movements of the
trade winds (i.e., summer Guinea Current influence versus winter Canary
Current influence; Fig. 1) associated with the year-round upwelling
occurrence results in strong oceanographic instability and in annual
variations in sea-surface temperature of about 10uC in the Golfe
dArguin.
Variable and eutrophic conditions lead to a rather simple food web
(opportunistic species and generalist predators) based on phytoplankton

Temperate-polar carbonates

Eutrophic tropicalsubtropical
carbonates

Tropical carbonates

rare
bivalves, small benthic and
planktonic foraminifers,
barnacles, echinoderms,
serpulids; few calcareous red
algae
rare to abundant
poor to good
aragonite dominant
destructive (grain dissolution and
maceration, biodegradation)

as opposed to an oligotrophic food web based on benthic algae, detritus,

and symbioses (cf. Wood 1993; Taylor 1997; Mutti and Hallock 2003;
Pomar and Hallock 2008). Eutrophic conditions thus suppress phototrophic and mixotrophic (phototrophic or heterotrophic) benthic organisms that are typical warm-water-related carbonate producers (i.e.,
hermatypic corals, calcareous green algae, large foraminifers). In
consequence, an interpretation of carbonate associations as tropical versus
extratropical environments needs to include consideration of the trophic
conditions. At a shelf scale, a heterozoan carbonate association overwhelmingly dominated by suspension and deposit feeders (e.g., infaunal
bivalves, but also sponges) indicates an ecosystem of highly productive
environment. In this eutrophic, aphotic setting, latitude and/or climatic
paleointerpretation is based upon the recognition of taxa (e.g., tropicalrelated mollusks) with well-constrained paleogeographic distribution.

FIG. 8.Distribution of the main skeletal

grains on the shelf of the Golfe dArguin and
factors controlling the neritic
carbonate production.

654

J. MICHEL ET AL.
CONCLUSIONS

The modern sedimentary system presented here is unique in the modern

world, in that it is a eutrophic tropical carbonate depositional system.
Nutrient concentration was previously shown to be an important
parameter of carbonate production (e.g., Hallock and Schlager 1986;
Halfar et al. 2004); however, so far no tropical carbonate association has
been presented that is overwhelmingly dominated by bivalves. This case
study presents a faunal assemblage characteristic of a eutrophic
environment, namely suspension-feeder-dominated carbonate grain
associations. This demonstrates that the perception of temperaturerelated carbonate sedimentation models (tropical vs cool-water carbonates) is not sufficient. This interpretation has strong implications for
paleolatitude and paleoclimatic studies.
ACKNOWLEDGMENTS

Thanks are due to the crews and scientists of the R/V Poseidon-346 and R/
V Poseidon-366 cruises and to the authorities of Mauritania for permission to
work in their exclusive economic zone. Karl Gurs, Serge Gofas, and Rudo
von Cosel are thanked for sharing their expertise on mollusk taxonomy. Karl
Gurs did not live to see this study completed. He is sorely missed. We are
grateful to Nereo Preto, John Reijmer, and Ralph Batzel for their support.
This paper greatly benefited from the reviews by Werner Piller, Cam Nelson,
an anonymous reviewer, AE Elias Samankassou, JSR editor Gene Rankey,
and CE John E. Southard. The MARUM (DFG-Research Center/Excellence
Cluster The Ocean in the Earth System) is acknowledged for providing
infrastructure and support for this research. This study was supported by the
German Science Foundation (We-2492/5). Two appendices are available from
JSR Data Archive, URL: http://sepm.org/pages.aspx?pageid5229.
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Received 16 April 2010; accepted 6 April 2011.