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Continental Shelf Research

journal homepage: www.elsevier.com/locate/csr

Western South Atlantic seasonal variability recorded in a mid-deglacial

bivalve from the outer Uruguayan continental shelf
André Klicpera a, b, h, Alvar Carranza c, *, Cristiano M. Chiessi d, Paula Franco-Fraguas e,
Till J.J. Hanebuth f, Hildegard Westphal a, b, g
a
Leibniz Centre for Tropical Marine Research (ZMT), Bremen, Germany
b
Department of Geosciences, MARUM, University of Bremen, Bremen, Germany
c
Departamento de Ecología y Gestión Ambiental, Centro Universitario Regional Este (CURE), Universidad de la República, Maldonado, Uruguay
d
School of Arts, Sciences and Humanities, University of São Paulo (USP), São Paulo, Brazil
e
Ciencia y Tecnología Marina (CINCYTEMA), Universidad de la República, Montevideo, Uruguay
f
Department of Marine Science, Coastal Carolina University (CCU), Conway, SC, USA
g
Physical Science and Engineering Division, King Abdullah University of Science and Technology (KAUST), Thuwal, Kingdom of Saudi Arabia
h
Área Biodiversidad y Conservación, Museo Nacional de Historia Natural, Montevideo, Uruguay

A R T I C L E I N F O A B S T R A C T

Keywords: The oceanographic dynamics on the continental shelf off southeastern South America are primarily controlled by
Retrotapes exalbidus the southward-flowing warm Brazil Current, converging with the northward-directed cold Malvinas (Falkland)
Brazil-Malvinas Confluence Current, and interacting with the continental discharge of the Plata River. The seasonally reversing regional wind
Subtropical shelf front
field together with the seasonal cycle of riverine discharge, determines which of these three components provides
Río de la plata estuary
Deglacial
the dominant forcing. The Uruguayan shelf is thus located in a transitional zone that extends from the region
influenced by the Brazil-Malvinas Confluence (BMC) in the open ocean to the Subtropical Shelf Front (STSF) on
the continental shelf. Understanding how the resulting oceanic seasonal variability responded to different cli­
matic boundary conditions may shed light on its future behavior. This study presents the first reconstruction of
mid-deglacial seasonal hydrographic variability on the continental shelf off southeastern South America in
seasonal resolution based on stable oxygen and carbon isotopes (δ18O, δ13C) from a thick-walled shell of a long-
living bivalve. The mid-deglacial (14.3 cal ka BP; Bølling-Allerød interstadial) Retrotapes exalbidus bivalve shows
a mean δ18O of 3.27 ± 0.42‰ (2.50 ± 0.42‰ when corrected for changes in global ice volume) and a seasonal
δ18O amplitude of 1.69‰ for raw isotopic excursions. Moreover, the δ13C exhibits abrupt negative peaks coin­
cident with more negative δ18O values that indicate seasons of elevated freshwater discharge. Finally, the growth
rate of the bivalve suggests that the specimen was closer to the metabolically optimum than modern individual of
this species from southern South America. Combining biogeographic and ecologic information with these iso­
topic data, the results point to colder waters and a slightly lower mid-deglacial seasonal amplitude in temper­
ature compared with modern conditions at this shelf site. Because of the northward-displaced Plata River mouth
during deglacial times, negative δ13C peaks are expected to reflect an influence of non-point freshwater sources
in the form of small fluvial distributaries along the paleo-coast. Most of this signal may, however, be driven by
seasonal metabolic effects associated with low ambient water temperatures related to a shallow-water envi­
ronment located closer to the respective paleo-coastline due to the low sea level at those times.

1. Introduction zone, the Brazil-Malvinas (Falkland) Confluence (BMC), formed by the

convergence of the Malvinas (Falkland) Current (MC) and the Brazil
The southeastern South American (SESA) continental shelf offshore Current (BC) (Olson et al., 1988; Lumpkin and Garzoli, 2011); (b) the
northeastern Argentina and Uruguay (Fig. 1) is of particular interest for vast Plata River estuary (RdlP), which supplies freshwater from the
oceanographic studies because it hosts: (a) a major oceanic confluence second largest drainage system in South America (Piola et al., 2005;

* Corresponding author..
E-mail address: alvardoc@fcien.edu.uy (A. Carranza).

https://doi.org/10.1016/j.csr.2023.105014
Received 1 February 2023; Received in revised form 25 April 2023; Accepted 7 May 2023
A. Klicpera et al.
Campos et al., 2008); and (c) a dynamic atmospheric system that is
influenced by the South American monsoon and seasonally reversing
wind fields with the dominance of southwestern winds in the austral
winter, and of southeastern winds in the austral summer (Matano et al.,
1993; Lumpkin and Garzoli, 2011).
The seasonal latitudinal and offshore displacement of shelf waters is
traceable through physico-chemical seawater properties along the con­
tinental shelf and the associated ecosystems (Piola et al., 2018, and
references therein). Recent studies suggested that the BMC, including its
shallow-water extension in the form of a vertical hydrographic boundary
known as the Subtropical Shelf Front (STSF), has shifted southwards
over the past century and particularly over the past few decades as a
response to current climatic change (Lumpkin and Garzoli, 2011; Bender
et al., 2013; Combes and Matano, 2014; de Souza et al., 2019). Under­
standing how this seasonal variability changed under different boundary
conditions may shed light on its future behavior. Therefore,
seasonally-resolved studies comparing the modern situation with past
conditions are instructive. Previous paleoceanographic studies from the
western South Atlantic reconstructed changes in the BMC and the STSF
with millennial-scale resolution (Bender et al., 2013; Lantzsch et al.,
2014; Voigt et al., 2015; Boretto et al., 2020; Gu et al., 2019), but the
seasonal variability remained elusive, mainly due to the lack of appro­
priate archives.
Sclerochronological archives from large bivalve species known for
their long individual life spans exceeding several decades provide
extremely valuable paleoceanographic information based on high-
resolution analyses (e.g., Dettman and Lohmann, 1995; Jones and
Quitmyer, 1996; Ivany and Runnegar, 2010; Lomovasky et al., 2002b;
Schöne et al., 2005). Here, we micro-sampled the multi-annual shell
increment succession of a fossil specimen of Retrotapes exalbidus, a
long-living marine bivalve. We performed δ13C and δ18O analyzes in
sub-seasonal to daily resolution, in order to gain insights into the dy­
namics of: (1) the intra and inter-annual variability in oceanographic
conditions; (2) the latitudinal displacement of the STSF driven by the
competition between the BC and the MC; and (3) the intensity of the
Plata River freshwater signal related to the changing rainfall intensity
over SESA. The new data provide key information for understanding the
long-term regional climate and ocean circulation over the continental
shelf. Finally, the isotopic information stored in the shell was compared
with the observed patterns in modern oceanographic stations along the
study region.
2. Study area and settings
The Uruguayan continental shelf is more than 150 km wide and hosts
a mosaic of markedly diverse submarine topographic elements and types
of sediment, where current-controlled coarse-grained shelf deposits may
occur in close vicinity to relief-confined mud depocenters (Urien et al.,
1980; Violante and Parker, 2004; Brazeiro et al., 2003). The inner shelf
is dominated by a bathymetric depression with late Pleistocene sedi­
ments (Fig. 1b), and a zone of partly carbonate-rich sand ridges adjacent
to this depression (Lantzsch et al., 2014). Numerous local autochtho­
nous and par-autochthonous shell beds are found on the mid and outer
shelf that consist of fine-to coarse-grained calcareous detritus (Violante
and Parker, 2004). Towards the shelf break, large areas of the shelf are
polished by strong confluence-related bottom currents, and deposition is
restricted to confined locations (Violante and Parker, 2004; Matano
et al., 2010).
From an oceanographic perspective, the SESA margin is currently
characterized by the confluence of the warm and saline BC (>20 ◦ C,
>36‰) flowing from the North, and the colder and less saline MC
(<15 ◦ C, <34‰) from the South (Fig. 1). These two currents converge
between 36 and 40◦ S, where they form the BMC (Fig. 1; Piola et al.,
2018, and references therein).
The STSF is a marked vertical hydrographic boundary located at
50–100 m water depth offshore Uruguay (32◦ S to 36◦ S), which separates
the colder Subantarctic Shelf Water (SASW, <11 ◦ C, <34.5‰), a branch
of the MC, from the warm and more saline Subtropical Shelf Water
(STSW, >16 ◦ C, >34.8‰) carried by the BC (Brandini et al., 2000; Piola
et al., 2000, 2008). Both the STSF and the BMC migrate latitudinally by
about 3◦ due to the reversing wind regime, reaching northern locations
in the austral winter, and southern ones in the austral summer (Schmid
and Garzoli, 2009). The shelf surface water (<20 m water depth) un­
dergoes large seasonal fluctuations in sea surface temperature (SST)
with amplitudes of about 10 ◦ C at 33◦ S and 3 ◦ C at 23◦ S (Piola et al.,
2000). Sea-surface salinity (SSS) shows only minimal seasonal vari­
ability (33.6–33.7‰) south of 37◦ S. Between 37◦ S and 35◦ S, in contrast,
a significant surface water freshening (<25–30.0‰) is related to the
large Plata River plume (Emilsson, 1961; Piola et al., 2000). This low
salinity plume is transported northwards close to the coast by the Bra­
zilian Coastal Current (BCC) (de Souza and Robinson, 2004).
The Plata River is fed by the Paraná and Uruguay Rivers covering
20% (3.2*106 km2) of the South American continent (Campos et al.,
2008). The mean discharge of the Paraná River (over the years
Fig. 1. Modern schematic oceanographic setting of
the study area on the southeastern South American
continental margin and location of the sediment core
taken on the outer shelf (A). The glacial paleo-valley
of the Plata River (white dashed arrow) (Lantzsch
et al., 2014) during the shelf exposure and the hy­
pothesized northward displacement of the
Brazil-Malvinas Confluence are illustrated in (B).
RdlP– Rio de la Plata plume BC – Brazil Current; BMC
– Brazil-Malvinas Confluence; SASW – Subantarctic
Shelf Water; STSF – Subtropical Shelf Front; STSW –
Subtropical Shelf Water.
A. Klicpera et al.

1884–1975) is 17,000 m3s-1 with a maximum of 22,000 and a minimum
of 8000 m3s-1. The mean discharge of the Uruguay River (1916–1975) is
4700 m3s-1 with a maximum of 14,300 m3s-1 and a minimum of 800
m3s-1. The discharge of these rivers peaks at different times during the
year, resulting in only minor seasonal variability in the Plata River
discharge to the ocean overall (5% of the annual mean; Burrage et al.,
2008). The present-day annual mean Plata River discharge is about 25,
000 m3s-1 (Wells and Daborn, 1997; Piola et al., 2018, and references
therein) and leads to the formation of the Plata Plume Water (PPW;
salinity <31‰; Depetris and Griffin, 1968). The PPW detaches from the
seafloor on the inner Uruguayan shelf and forms a major, coast-parallel
buoyant tongue transported northwards by the BCC (Fig. 2; Wells and
Daborn, 1997). Conductivity and temperature through depth (CTD) data
(Antonov et al., 2010; Locarnini et al., 2010) indicate that the PPW
occupies the upper 10 m of the water column in austral summer
(January–March), and in the upper 20 m in austral winter (July–Sep­
tember), flowing along the inner Brazilian shelf as far north as 26◦ S
(Fig. 2; Piola et al., 2000).
This oceanographic setting controls the regional biogeographic pat­
terns observed in benthic mollusks. Currently, the Uruguayan shelf hosts

Fig. 2. Modern oceanographic conditions off southeastern South America for austral summer (A, B, E, F) and austral winter (C, D, G, H). Salinity (A to D) and
temperature (E to H) are shown for both seasons. The latitudinal shelf sections shown in panels A, C, E and G (left column) follow a southeast-northeast transect
marked as dashed line in panel B. SASW – Subantarctic Shelf Water; PPW –Plata Plume Water; STSW – Subtropical Shelf Wate; STSF – Subtropical Shelf Front. Data:
World Ocean Atlas 2010 (Antonov et al., 2010; Locarnini et al., 2010).
A. Klicpera et al.
a malaco-faunal transition zone between the magellanic (to the South)
and subtropical (to the North) benthic biogeographic provinces (Olivier
and Scarabino, 1972; Kaiser, 1977; Floeter and Soares-Gomes, 1999;
Scarabino, 2003). In addition, a separate benthic mollusk assemblage
occurs along the inner shelf under the influence of the Plata River, where
its diversity strongly correlates with the salinity gradient. The outer shelf
hosts both magellanic and subtropical elements, and benthic species
turnover is associated with latitude (Carranza et al., 2008). The steep
meridional temperature gradient that characterizes both BMC and STSF
may act as a biogeographic barrier preventing an exchange between
Antarctic cold-water and subtropical warm-water species, as evidenced
for at least eight cold-water bivalves exhibiting their northernmost
distribution boundary in the area; associated with offshore sub-Antarctic
waters (Scarabino et al., 2015). Thus the occurrence of a given fossil or
subfossil mollusk species can be highly informative of the biogeographic
setting in which they lived, informing about the paleo-distribution of
water masses in the study area.
3. Material and methods
3.1. Bivalve shell and taxonomic identification
During research cruise M78 Leg 3a performed with the German R/V
METEOR in 2009 (Krastel et al., 2012), a large, articulated, intact
bivalve shell was recovered from the base of Vibracore GeoB13802-2
(345 cm recovery; bivalve sampling depth at 335–341 cm), taken near
the shelf break at 141 m water depth (Fig. 1b; Table 1). The bivalve
specimen was identified to species level as Retrotapes exalbidus (Dillwyn,
1817) aka Eurhomalea exalbida (Dillwyn, 1817) (the “Ring Clam”)
through comparison with reference material housed at the Museo
Nacional de Historia Natural (MNHN), Montevideo, Uruguay (deter­
mined by F. Scarabino). R. exalbidus is a cold-water species that lives
within the shallow sub-littoral zone. With a maximum life span of about
70 years and massive valves that reach up to 9 cm length, this bivalve is
considered a useful climate archive (Gordillo et al., 2015). Species
nomenclature used in this study is based on MolluscaBase Editorial
Board (2022). The description of the inner bivalve structures (e.g.,
shell-layer configuration) follows th e revised terminology of Schöne
et al. (2011).
3.2. Microscopy of shell preservation status and shell microstructures
The suitability of a specimen for isotopic analysis depends on its
preservation status. Possible impacts by bioerosion, recrystallization, or
physical abrasion were assessed visually in petrographic thin sections
using reflected and transmitted light microscopy. Thin sections were
oriented along the maximum growth axis of the shell valve and groun­
ded to a slide thickness of 50 μm. The thin sections were stained with
Feigl’s Solution to confirm its original aragonitic mineralogy (Leitmeier
and Feigl, 1934). Growth increment width was measured on the petro­
graphic thin sections across the outer shell (Fig. 3a) and the hinge sec­
tion (chondrophore; Fig. 4b) using the image analysis software ImageJ
1.45s. The microstructure was investigated with a Tescan Vega 3 XMU
scanning electron microscope (SEM, 20.0 kV, secondary electron mode)
at the Leibniz Center for Tropical Marine Research (ZMT), Bremen,
Germany. Samples for SEM analysis were cut from those umbonal and
central shell positions (Fig. 3a), where growth lines and microstructures
were found in well-preserved conditions. The samples were etched with
hydrochloric acid (0.25% HCl) for 10–20 s to better reveal the growth
lines, and gold-sputtered afterwards.
Table 1
Sampling location. GeoB – Bremen/MARUM Core Repository; VC – vibracore.
Sediment core Latitude (◦ S) Longitude (◦ W) Coring gear
GeoB13802-2 36 05.30
◦ ′
53 20.72
◦ ′
VC
3.3. Carbonate mineralogy of inner and outer shell layers
The mineralogy of shell carbonate powder samples was determined
at the Central Laboratory for Crystallography and Applied Material
Sciences (ZEKAM, University of Bremen) by means of X-ray diffraction
(XRD) (Fig. 3a). A Philips X’Pert Pro multipurpose diffractometer
equipped with a Cu-tube (k" 1.541, 45 kV, 40 mA), a secondary Ni-Filter
and an X’Celerator detector system were used for continuous scans from
3 to 85◦ 2θ with a calculated step size of 0.016◦ 2θ. Minerals were
identified with the Philips software X’Pert HighScore and the X-ray
diffraction interpretation software MacDiff version 4.25.
3.4. Radiocarbon dating
The shell was sampled for accelerator mass spectrometry radio­
carbon (AMS-14C) dating in two positions, namely the juvenile umbonal
shell (dorsal) and near the shell margin (ventral) (Fig. 3b, d). Powder
samples were micro-drilled along marginal transects of 10 mm in total
length, set parallel to the growth bands. The resulting samples of 30–50
mg comprised less than two growth increments. AMS-14C dating was
carried out at the Poznan Radiocarbon Laboratory, Poland. The age of
the R. exalbidus specimen was calibrated with the CALIB v. 8.2 software
and the Marine20 dataset, using the automatically included conserva­
tive global reservoir age of 550 years (Stuiver et al., 2020; Heaton et al.,
2020).
3.5. Stable oxygen and carbon isotope analyses
A MicroMill system (New Wave Research) was used for high-
resolution sampling, which is designed to recover 40–50 μg of powder
for isotopic analysis (Dettman and Lohmann, 1995). Only
well-preserved shell areas were drilled while altered areas were omitted.
For sampling the shells, 50-μm thin sections were cut and polished.
Horizontal drilling speed was adjusted to 700 μm/s, milled in eight
consecutive passes of 35 μm depth. The sampling track distance was 100
μm. Digital images of the shell sections were taken with a Leica DFC-280
digital camera attached to a Leica binocular microscope (Fig. 3b). The
bivalve was sampled along a 2.25 cm long transect encompassing six
growth increments (AI-1 to AI-6) with 256 single tracks (T1 to T256;
Fig. 3B). For obtaining sufficient carbonate material, up to three drilling
tracks in the outer shell layer were combined in one sample, resulting in
a sampling resolution of more than 10 samples per year and 66 samples
for stable isotope analysis (Fig. 3e: RE1 to RE66). Additional samples
were drilled from the inner shell layer (n = 23) placed in increment I–C
(Fig. 3e: REA1 to REA13) and increment I-D (Fig. 3e: REA14 to REA23).
The powder micro-samples were analyzed for δ13Cshell and δ18Oshell
using a Kiel III automated carbonate preparation device connected on­
line to a ThermoFinnigan MAT 252 isotope-ratio mass-spectrometer
coupled to a GasBench II (Kim et al., 2007). All isotopic values are re­
ported in the conventional δ-notation in parts per mil (‰) relative to the
Vienna Pee Dee Belemnite (VPDB) using the NBS19 standard (δ18O =
− 2.20‰). Reproducibility was evaluated by replicate analysis of labo­
ratorial standards to ±0.03‰ 1σ for δ13Cshell and ±0.05‰ 1σ for
δ18Oshell.
3.6. Modern environmental data integration
To capture the potential influence of both continental water
discharge and alternating shelf water masses on the stable isotopic sig­
nals recorded in the R. exalbidus shell, we obtained the modern oceanic
Water depth (m) Recovery (cm) Species
141 341 Retrotapes exalbidus
A. Klicpera et al.
Fig. 4. (A) Shell height versus age plot for the investigated deglacial Retrotapes
exalbidus specimen (black line) in comparison to previously reported modern
specimen growth records (grey area; Lomovasky et al., 2002a) indicates a later
onset of maturity for the investigated specimen. (B) Full growth record of the
chondrophore (hinge section) of the investigated Retrotapes exalbidus is pre­
sented as a mosaic of ten thin sections.
conditions (temperature, salinity), and estimated both stable oxygen
isotopic composition of seawater (δ18Osw) and corresponding δ18O
values precipitated in equilibrium (δ18Oeq) at virtual stations located
along inner and outer shelf. The selected five “coastal” stations on the
inner shelf (C1 to C5) were distributed close to the 50 m isobath, and five
outer shelf stations (O1 to O4) located close to the 150 m isobaths,
including the particular site where the R. exalbidus shell was recovered
(E Core) (Fig. 7).
Fig. 3. (A) Photo-micrograph of the investigated
deglacial specimen of Retrotapes exalbidus, (B) Close-
up of growth increments sampled (AI-1 to AI-6) and
MicroMill transects T1 to T256 on the outer shell; (C)
Schematic illustration of sampling scheme. Accre­
tionary increments were sampled starting at an age of
nine years onward in order to avoid metabolically
driven fractionation effects during early ontogeny
(AI-1 to AI-6 in panels B and C). (D) At a higher
resolution, each annual growth increment provides a
record of several (monthly) sub-increments, which
(E) at highest resolution show daily micro-growth
increments of 30–80 μm in individual width. ISL –
inner shell layer; OSL – outer shell layer; iOSL –
innermost outer shell layer; oOSL – outermost outer
shell layer.
We used averaged monthly temperature and salinity data from the
World Ocean Atlas 2010(Antonov et al., 2010; Locarnini et al., 2010) for
water masses identification (subtropical and subantartic water masses)
and an estimation of δ18Osw and δ18Oeq. For the estimation of δ18Osw
(VSMOW) based on salinity, the Tropical Atlantic (Equation (1);
LeGrande and Schmidt, 2006) and the South Atlantic (Equation (2);
LeGrande and Schmidt, 2006) equations were applied concerning the
subtropical and subantarctic water masses, respectively. An alternative
equation, calibrated for the shallow shelf and related to subantarctic
waters (Equation (3); Colonese et al., 2012), was used for comparison
reasons. A paleo-temperature equation for aragonite (Equation (4);
Grossman and Ku, 1986) was used for estimating δ18Oeq (VPDB).
Δ18Osw = 0.15 * S – 4.61 (1)
Δ18Osw = 0.51 * S – 17.4 (2)
18
Δ Osw = 0.28 * S – 10.79 (3)
◦ 18 18
T( C) = 20.6–4.34 * (δ Oeq – (δ Osw (SMOW) – 0.27)) (4)
18
The seasonal range (amplitude) in δ Oeq estimated for each virtual
oceanographic station was compared with the ice volume-corrected
δ18Oivc seasonal range of the R. exalbidus fossil specimen (δ18Oivc-shell).
During deglacial low-stand sea level conditions (Guilderson et al., 2000;
Lambeck and Chappell, 2001) δ18Osw was ca. 0.77‰ (0.0081‰/m,
Fairbanks and Matthews, 1978; Lambeck and Chappell, 2001) higher
than modern δ18Osw. Hence, 0.77‰ was extracted from δ18Oshell before
comparison with estimated δ18Oeq. Since there is to our knowledge no
species-specific paleotemperature equation for R. exalbidus we are
A. Klicpera et al.
unable to directly compare absolute δ18Oeq values to δ18Oivc-shell values.
However, because the slopes of different paleotemperature equations
are similar (e.g., Grossman and Ku, 1986; Böhm et al., 2000; Kim et al.,
2007) we can compare the seasonal range of the δ18Oeq to the seasonal
range of the δ18Oivc-shell.
3.7. Temporal alignment of Retrotapes exalbidus δ18Oshell
In order to compare the estimated δ18Oeq time series obtained from
modern stations that are provided as monthly values together with the
δ18Oivc-shell time series, both data series have to be plotted against the
same time axis. Importantly, modern WOA09 (Antonov et al., 2010;
Locarnini et al., 2010) data are monthly averaged over several decades,
whereas each shell value analyzed here represents those environmental
conditions that prevailed during a specific month or a short series of
month. Thus, shell raw data are assumed to record absolute values,
while modern data are smoothed, showing average conditions and, thus,
underrepresenting the true amplitude. To reduce this bias, we deter­
mined the average minimum and maximum of δ18Oeq within an annual
cycle. We found that δ18Oeq maxima, averaged across the region, occur
in September and minima in April. Then, we assigned the maximum
δ18Oivc-shell value within each annual increment (distinct micro­
structurally units that form over a period of several months each year) to
match with September values in the δ18Oeq time series and, conversely,
the minimum δ18Oivc-shell to match April in the δ18Oeq time series.
Missing values in the δ18Oivc-shell time series were reconstructed using a
moving average between consecutive values of δ18Oivc-shell. Thus, we
obtained a monthly time series for the δ18Oivc-shell that allowed us to
compare δ18Oeq and δ18Oivc-shell in equal statistical footing (i.e., 12
values each per year; averaging monthly values for the six annual in­
crements). This procedure reduces the variability of δ18Oivc-shell. Further,
one way Analysis of Variance (ANOVA) was performed to evaluate
differences in mean values between the six annual increments sampled
in the analyzed R. exalbidus specimen. Significant differences may
indicate the differential action of vital effects during the observed
interval.
4. Results
4.1. Species determination, shell growth patterns, mineralogy and
preservation status
The articulated shell from the base of core GeoB13802-2 (Fig. 1;
Table 1) was identified as Retrotapes exalbidus (Dillwyn, 1817). The
R. exalbidus specimen shows a well-defined shell-layer configuration
with a well-preserved inner shell layer (ISL; Fig. 3a). The shell in­
crements are devoid of bioerosional traces and exhibit a very distinct
growth-line pattern within the hinge plate that forms the chondrophore.
The ontogenetically youngest/juvenile increments illustrate high calci­
fication rates (rapid shell growth) with a pronounced decrease in
increment thickness towards ontogenetically older/adult increments.
SEM imagery revealed well-preserved, composite prismatic structures
(aragonite) for the outer shell layer (oOSL), subdivided by finer growth
lines that run parallel to the main growth lines (possibly daily growth
increments; Chateigner, 2000; Gobac et al., 2009). XRD analysis
revealed an aragonite-dominated mineralogy (Fig. 3e) with an inner
shell layer (iOSL) of purely aragonitic nature and an oOSL composed of
92% aragonite (8% calcite). Both oOSL and iOSL provided an excellent
quality record for sampling in the highest resolution.
Light microscopy revealed 25 paired major first-order growth in­
crements (dark band followed by a filigree white 50–100 μm thick
growth line), subdivided by (sub-annual) second-order growth in­
crements, a pattern that was consistent along the entire oOSL transect
(Fig. 3a). The same pattern was observed as a compressed record in the
chondrophore section (Fig. 4b) and the iOSL (Fig. 3b, d). The width of
the first-order growth increments found in the analyzed specimen
throughout the shell record ranged from 23.11 mm for the earliest ju­
venile increment to 0.23 mm for the latest adult increment, with an
average increment width of 4.03 mm (N = 25).
Second-order subordinate increments (herein called micro-
increments) were obvious in the R. exalbidus shell as well (Fig. 3c, d,
e). In the collected specimen, the second-order increment widths ranged
from 9 to 164 μm (average = 46 μm), being narrowest towards the
annual growth lines and widest in the center of a first-order increment.
The analysis of the first 20 years, i.e., during juvenile growth gives a
mean growth rate of 3.96 mm yr− 1 (10.85 μm day− 1).
4.2. Age control and stable oxygen and carbon isotopes micro-transects
The R. exalbidus specimen was dated around a median average age of
14,275 cal ka BP (Table 2; Fig. 3d). This radiocarbon age corresponds to
mid-deglacial times and coincides with the Bølling-Allerød interstadial
and the end of Meltwater Pulse 1A (Hanebuth et al., 2000). The
high-resolution δ18Oshell profile (N = 66, Fig. 5a) from the oOSL of
R. exalbidus reveals a mean δ18Oshell(oOSL) value of 3.27 ± 0.42‰, and,
after correction for ice volume, a value of 2.50 ± 0.42‰ (data provided
in the supplementary table). The uncorrected δ18Oshell(oOSL) amplitude is
1.69‰ with a maximum value of 3.98‰ and a minimum of 2.29‰. The
temporally unaligned δ18Oshell(oOSL) signature (Fig. 5c; Suppl. Table 1)
exhibits successive cycles of rapidly increasing values occurring shortly
after growth-line formation, and more gradually decreasing values until
the newt growth line. As a result, the δ18Oshell(oOSL) signature describes a
characteristic asymmetric cyclic pattern. The time series shows most
negative δ18Oshell(oOSL) values (high water temperatures) associated with
annual growth lines, while the most positive δ18Oshell(oOSL) values (low
water temperatures) appears shortly after the first third of growth in­
terval (Fig. 5a). No significant differences were found between the mean
values of the six annual increments (ANOVA; p > 0.05), and thus no
significant ontogenetic trend was found (see Table 3).
The δ13Cshell(oOSL) signal shows a mean value of 0.41 ± 0.49‰. The
13
δ Cshell(oOSL) amplitude range is 2.76‰ with a maximum of 1.02‰ and
a minimum of − 1.74‰. The unaligned δ13Cshell(oOSL) signature exhibits a
slight trend towards lighter values per annual growth interval, inter­
rupted by sharp cyclic minima, being increasingly pronounced with
ontogeny. These negative excursions are closely associated with the five
growth lines (Fig. 5c) and thus indicate a distinct positive correlation
with δ18Oshell(oOSL) minimum values.
4.3. Environmental setting
The investigated R. exalbidus specimen lived during mid-deglacial
times (14.5 cal ka BP). Although the regional post-LGM sea-level his­
tory is not yet well-understood (Violante and Parker, 2004, and refer­
ences therein), the eustatic sea-level at 14.5 cal ka BP can be roughly
estimated to a position at or lower than 110 m below present-day
sea-level (Lambeck and Chappell, 2001; Hanebuth et al., 2009). Given
that the articulated R. exalbidus studied here was found at 145 m modern
water depth (considering the sample depth below modern seafloor), its
living depth can, accordingly, be estimated to a paleo-water depth of not
more than 50 m, i.e., the nearshore to inner-shelf zone. This reduced
depth is in agreement with Rosenberg (2009) and Häussermann and
Försterra (2009) who reported modern depth ranges for this species as
maximum depths of 70 and 150 m.
4.4. Modern conditions
In general, all stations present seasonal and salinity temperature
oscillations with higher values in austral summer/autumn and lower
ones in austral winter/spring. Salinity shows relatively less pronounced
seasonal oscillations than temperature (Suppl. Fig. 1). The virtual sta­
tions south of 37◦ 30′ S (Stations O1, O2, C1, C2) show reduced ampli­
tudes in δ18Oeq compared with the stations further north (Fig. 6). On the
A. Klicpera et al.

Table 2
Accelerator mass spectrometry 14C ages of dorsal (ontogenetic youngest) and ventral (ontogenetic oldest) samples from the investigated Retrotapes exalbidus specimen
in sediment core GeoB13802-2. GeoB – Bremen/MARUM Core Repository; POZ – Poznan Radiocarbon Laboratory, Poland.
14
Lab ID Species Depth in core (cm) Position in shell Raw C age (yr BP) 1σ age range (cal yr BP) Median age (cal a BP)

Poz-36091 R. exalbidus 341 dorsal 14,200 ± 70 14,340–14,080 14,340

Poz-36092 R. exalbidus 341 ventral 14,060 ± 60 14,530 - 14,250 14,210

Table 3
Statistical parameters of the stable carbon (δ13Cshell) and oxygen (δ18Oshell) isotopic records from the investigated deglacial Retrotapes exalbidus shell. (*) Location of
micro-sample in across-shell profile: oOS – outermost outer shell layer; iOSL – innermost outer shell layer. Ampl.: Total amplitude of isotopic range.
Species Location (*) δ18O (‰, VPDB) δ13C (‰, VPDB)

Range Mean Ampl. Range Mean Ampl.

R. exalbidus oOSL 2.29–3.98 3.27 ± 0.42 1.69 − 1.74–1.02 0.41 ± 0.49 2.76
R. exalbidus iOSL 2.93–3.97 3.44 ± 0.30 1.04 0.25–1.62 1.17 ± 0.37 1.37

Fig. 6. Comparison of the amplitude of the stable oxygen isotopic values

Fig. 5. (A and B) Schematic drawing of the investigated Retrotapes exalbidus
specimen showing annual growth intervals (AI) along the analyzed transect. (C)
Ice volume corrected stable oxygen isotopic values (δ18Oivc-shell) (continuous
line) and stable carbon isotopic values (δ13Cshell) (dashed line) of the investi­
gated R. exalbidus shell.
outer shelf, δ18Osw mean values increase towards the north from − 0.1‰
(O1) to 0.8‰ (O4). Mean δ18Oeq values decrease towards the north from
ca. 2.9‰ (O1) to 0.7‰ (O4). Comparisons of normalized amplitudes of
the δ18Oeq values for all virtual stations are shown in Fig. 6.
5. Discussion
5.1. Biogeographic evidence
In general, our findings are in agreement with the known paleo-
biogeographic distribution of faunal elements suggesting colder tem­
peratures during deglacial times off Uruguay. For example, Aguirre et al.
(2009, 2011) reported that faunal-associations showed typical
cold-water elements north of their present-day northernmost boundary
prior to the Holocene climatic amelioration, locally coinciding with the
BMC (38◦ S) offshore northern Argentina, linked to a stronger Malvinas
current. This trend reversed at least for the shallower ocean towards the
(δ18Oeq) calculated for modern conditions at the virtual stations (please see the
location of the virtual stations in Fig. 7) and monthly averages of the ice volume
corrected δ18Oshell of the analyzed specimen of Retrotapes exalbidus. Standard­
ized data show the relative amplitude of the isotopic signature for inner (C1 to
5) and outer (O1 to O4) shelf stations including the R. exalbidus core site (R
Core). Data are sorted from left to right based on minimum values.
Holocene when subtropical species even approached the Uruguayan
coast, suggesting the oceanic conditions have been warmer than at
present-day (e.g. Martínez et al., 2006; Aguirre et al., 2011).
The neoaustral genus Retrotapes first appeared in Patagonia and
Antarctica during early Tertiary times and has been described in
Miocene and Holocene records from southern South America (del Rio,
1997; Gordillo, 2006). R. exalbidus is extant in the Magellan Region
(Carcelles, 1944; Reid and Osorio, 2000). Its modern northernmost
biogeographic occurrence (ca. 38◦ S; Penchaszadeh et al., 2008) co­
incides with the northernmost position of the MC and the associated
SASW (Figs. 1 and 2). The investigated specimen was recovered at 36◦ S,
i.e., about 220 km north of the modern species distribution limit (38◦ S)
reported by Penchaszadeh et al. (2008). Scarabino et al. (2015), in
contrast, placed the modern northern boundary of R. exalbidus appear­
ance onto the Uruguayan shelf (North of 38◦ S)based on single detached
but freshly preserved valves that still contained their organic ligament.
These results may point to modern habitat conditions that might be
marginally suitable for such species in the Uruguayan shelf. This
observation is also congruent with inferred cooler paleoceanographic
conditions in the region during the Bølling-Allerød interstadial and the
end of Meltwater Pulse 1A and an inner-shelf environment with water
depth of (Guilderson et al., 2000; Hanebuth et al., 2000) during this time
A. Klicpera et al.
Fig. 7. Location of modern (virtual) stations along the study area. Retrotapes
exalbidus specimen was found in station R CORE.
at the study site.
5.2. Sclerochronology
The internal banding of the early deglacial R. exalbidus specimen
provided a record of 25 major growth bands that documents the growth
over a lifespan of 22–25 years. For comparison, R. exalbidus from the
Magellan region is known to live up to 70 years (Gordillo, 2006; Gordillo
et al., 2013a; Lomovasky et al., 2002b; Yan et al., 2012). Additionally,
the interpretation of annually precipitated increments in our data set is
corroborated by the stable isotopic signature, which provides
well-defined cycles interpreted to be seasonal (Fig. 5). This finding is
consistent with previous the interpretation of first-order increments as
representing annual growth pattern (Lomovasky et al., 2002a; Ivany
et al., 2008, and references therein).
The annual growth pattern of modern R. exalbidus shells has been
addressed in a number of studies (Dextraze and Zinsmeister, 1987;
Lomovasky et al., 2002a, 2002b; Ivany et al., 2008). Yan et al. (2012)
have shown that growth lines in modern shells do not simply reflect a
winterly cold-water signature, but that other key paramters coupled to
primary production, such as food availability, may have an influence on
biochemical carbonate precipitation processes and, thus, shell growth
pattern. Contrary to the metabolism-controlled growth of first-order
increments, intra-annual growth structures are thought to be resulting
from regional environmental parameters that include year-specific
short-term variability in water temperature, food availability (primary
production) and environmental stress factors (e.g., influencing the
timing of the spawning season; Gordillo et al., 2013). Control factors
causing environmental stress such as seasonality in temperature, but
also the reproductive activity, have been shown to influence shell
growth, demonstrated, for instance, on the long-lived North Atlantic
cold-water Artica islandica and the Patagonian Zygochlamys patagonica
(Jones 1980; Morriconi et al., 2002; Elliot et al., 2003; Schöne et al.,
2003, 2005; Lomovasky et al., 2008). It is, thus, likely that such control
mechanisms may also have influenced the R. exalbidus specimen studied
here.
The trend towards lighter δ13Cshell during ontogenesis has been
observed in a number of studies (e.g., Keller et al., 2002; Gillikin et al.,
2007). Keller et al. (2002) attributed more negative δ13C values in older
shell parts of Chamelea gallina to the utilization of δ13CDIC derived from
pore waters associated with deeper burrowing into the seabed sediment.
Gillikin et al. (2007) correlated this isotopic trend with metabolic
changes during the organism’s life cycle. Yan et al. (2012) also noted
such a trend for R. exalbidus. The existence of an ontogenetically
controlled, fast-growing juvenile phase and a successive reduction
during the adult stage is typical for bivalves and assumed to be related to
the onset of sexual maturity (Gosling, 2003). Modern specimens of
E. exalbida show, for instance, an average growth rate not exceeding
2.50 mm yr− 1 (N = 214; Fig. 4a) and an onset of the adult stage after
15–20 years (Lomovasky et al., 2002a). The R. exalbidus specimen
studied here reveals a higher growth rate for the first 20 years (mean
3.96 mm yr− 1 = 10.85 μm day− 1), followed by a delayed onset of
maturity (25–30 year) compared to modern individuals collected in
Ushuaia Bay (Tierra del Fuego, southernmost South America; Lomova­
sky et al., 2002a). Such an ontogenetic offset of our R. exalbidus spec­
imen towards higher growth rates and later maturity can be attributed to
oceanic conditions closer to its environmental metabolic optimum (i.e.,
colder water, normal salinity, sufficient food supply) than it is the case
today in Usuahia Bay. This can be linked to an exposure to a higher
oceanic alkalinity and dissolved calcium concentration associated with
less marked estuarine conditions during mid deglacial times, when
compared with modern salinity ranges observed in Usuahia bay (Gor­
dillo, 2006). For modern Z. patagonica, growth rates have been shown to
decrease along a geographic gradient southward of 39◦ 30′ S, while
Uruguayan stocks located between 35◦ 50′ S and 36◦ 50′ S show
decreasing growth rates towards the northern limit of its geographical
range (Gutierrez and Defeo, 2005). From our data, the scenario of an
exposure to a fully marine environment is unlikely, according to the
δ13C values (see below). Thus, the sampling site may have been located
closer to the metabolic optimum of the species in comparison with
modern populations living in the southern limit of its distribution.
5.3. Regional paleoceanographic and climatic implications
This study presents the first subannually resolved δ18Oshell record
from the early deglacial off SESA. The δ18Oshell record show most
negative values associated with annual growth lines and negative
δ13Cshell peaks. This covariation between δ18O and δ13C values would be
expected under seawater-freshwater mixing (e.g., Wefer, 1985; Ingram
et al., 1996; Ivany et al., 2008), suggesting short-term increases in the
discharge of fluvial waters into the marine realm. There is no evidence of
increased rainfall coevally with minima in the δ18Oshell record, since
summer precipitation related to the South American monsoon system
most probably was not substantially increased during the
Bølling-Allerød interstadial (Razik et al., 2013; Gu et al., 2018).
According to Urien and Ottman (1971) and Lantzsch et al. (2014),
the wide SESA continental shelf was largely exposed and characterized
by coastal plains during deglacial times, while the Plata River followed a
steeply northward-directed morphological depression right off the
modern Uruguayan coastline and was largely isolated from our sampling
site (Fig. 1a). Because of the Plata River paleo-mouth having been
located at the latitude of the Rio Grande Cone off southern Brazil during
deglacial times (e.g., Chiessi et al., 2008; Lantzsch et al., 2014; Gu et al.,
2018), negative δ13C peaks should, thus, reflect the influence of
non-point freshwater sources in the form of small fluvial distributaries
along the paleo-coast (e.g., Violante et al., 2008).
The amplitude of the raw δ13C signal from our specimen is higher
(2.76‰) compared with data from Yan et al. (2012). These authors re­
ported an average amplitude of 0.73‰ (the highest amplitude recorded
from an individual was 1.63‰), based on data from four young (4–10
A. Klicpera et al.
years old) modern specimens of R. exalbidus sampled at 51◦ 40′ S (salin­
ities averaging 33.68 ± 0.12). This observation reinforces the hypoth­
esis of more pronounced variation in salinity at the site of collection of
our bivalve specimen during the mid-deglaciation. As expected, our
modern virtual oceanographic stations on the inner shelf reflect riverine
freshwater dilution of shelf water masses. This dilution effect is partic­
ularly evident in intermediate stations close to the Plata River mouth
and reflects the highest continental discharge in the austral winter/­
spring (Simionato et al., 2001; Moller et al., 2001).
However, according to our temporally aligned δ18Oshell data, the
seasonality of the environment experienced by the R. exalbidus specimen
differed only slightly from what is observed today in the sampling site,
with the mid-deglacial specimen probably exposed to a reduced intra-
annual variability (Fig. 6). Standardized data clearly shows a lat­
itudinal trend in modern amplitudes, with stations located south of
36◦ 30’ (C1, C2, O1 and O2) exhibiting narrower amplitudes compared
with the remaining stations. In this comparison, data from the core site
(R Core) showed a very similar amplitude to the isotopic signal recov­
ered from the shell. Further, modern data from our virtual stations
(Fig. 7) indicate that the northernmost stations are bathed by relatively
warm and salty water masses (i.e., STSW) while the southernmost sta­
tions represent relatively cold and fresh water masses (i.e., SASW). In­
termediate stations are located at the confluence of both shelf water
masses (i.e., near the STSF). Over the Uruguayan shelf, below 100 m
water depth, seasonal changes in water temperatures are driven by the
relative dominance of STSW over SASW in autumn, whereas spring is
characterized by a dominance of SASW (Ortega and Martínez, 2007). In
contrast, the general seasonal temperature oscillation observed within
water masses likely reflects the cooling of the upper layer during cold
seasons (Piola et al., 2000). Taken together, the evidence points out to a
more pronounced dominance of SASW over shallower coastal areas
during Bølling-Allerød interstadial.
6. Conclusions
Although we recognise the limited value of the information retrieved
from a single specimen, this study presents the first high-resolution
reconstruction of seasonal oceanic variability on the continental shelf
off southeastern South America (SESA) during mid-deglacial times. We
analyzed the 6-years spanning δ18O and δ13C stable isotope record of an
in-situ preserved, long-living bivalve specimen. Allowing for a compar­
ative approach, the modern bathymetric and biogeographic distribution
pattern is known and stable isotopic data exist for modern R. exalbidus,
although the modern environmental conditions experienced by inves­
tigated individual organisms may not be fully comparable to those
environmental conditions experienced by the collected fossil specimen
that lived about 14.5 ka ago.
An accurate paleoceanographic reconstruction of ocean water pa­
rameters in the absence of species-specific paleo-temperature equations
is a challenging approach. Our new data help to characterize the
participating processes that were once active on the SESA continental
shelf. These data pointed out to colder waters during deglacial condi­
tions, and δ13C suggests a transition to a more variable salinity regime in
nearshore areas during the Bølling-Allerød interstadial. The δ18O and
δ13C record from the R. exalbidus specimen presented here shows coin­
cident and abrupt negative peaks that indicate pulses of freshwater
discharge. Further, since both series shows coincident peaks, the tem­
perature dependence of isotopic fractionation processes cannot be
excluded as a driver of the observed pattern. Despite the challenge of
identifying the exact mechanism responsible for these negative δ13C
excursions, the existence of minor fluvial distributaries with their river
mouths located all along the coast might offer a plausible explanation
for diverting the isotopic signals. Since the shell collection site was
located in shallow waters and closer to the respective mid-deglacial
coastline, this hypothesis is worth exploring in the future. Impor­
tantly, the overall amplitude of the δ18Oivc-shell record is similar to the
amplitude of modern temperature conditions on the R Core site, sug­
gesting only slightly lower mid-deglacial seasonal amplitude in tem­
perature compared to present day conditions. Future research may focus
on (i) producing a species-specific paleo-temperature equation for R .
Exalbidus, and (ii) dating and analyzing fossil specimens found on the
Brazilian shelf ca. 34◦ S.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Acknowledgements
Chief Scientist Sebastian Krastel-Gudegast and the participants and
crew of the Meteor Cruise M78/3a are gratefully acknowledged for
collecting the sample material and for providing sample background
data. Sebastian Flotow (ZMT Bremen, Germany) is thanked for prepar­
ing the thin sections. Matthias López Correa (GeoZentrum Nordbayern,
Germany) is thanked for support with the MicroMill sampling. Michael
Joachimski, Daniele Lutz (both GeoZentrum Nordbayern, Germany)
performed stable oxygen and carbon isotopes analyses. We also thank
Felipe García-Rodríguez (Facultad de Ciencias, UdelaR Montevideo,
Uruguay) and Fabrizio Scarabino (Museo Nacional de Historia Natural
in Montevideo, Uruguay) for their local support and helpful discussions.
Leonardo Ortega is acknowledged for his help with the oceanographic
analyses. This project was funded through the DFG-Research Center/
Cluster of Excellence “The Ocean in the Earth System” Project SD-2. C.M.
C. acknowledges the financial support from FAPESP (grants 2018/
15123–4, and 2019/24349–9), CNP (grant 312458/2020–7), CAPES
(grant 88881.313535/201901) and the Alexander von Humboldt
Foundation.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.csr.2023.105014.
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