Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255

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Are productivity and strati¢cation important to

sapropel deposition? Microfossil evidence from
late Pliocene insolation cycle 180 at Vrica, Calabria
Alessandra Negri  , Caterina Morigi, Simona Giunta
Istituto di Scienze del Mare, Universita’a¤ di Ancona, Via Brecce Bianche, 60131 Ancona, Italy
Received 26 March 2002; received in revised form 26 June 2002; accepted 18 October 2002

Abstract

We present the results of a micropaleontological study performed on the sapropel sequence associated with
insolation cycle 180 from the Plio^Pleistocene Vrica sequence (Calabria, Italy). We performed a high-resolution study
on the 3.38-m-thick layer c from a core drilled close to the classical outcrop section in which we analyze fluctuations
in the abundance and composition of calcareous nannofossils and planktic and benthic foraminifera. Changes in the
fossil assemblages reveal at least three major paleoenvironmental phases in layer c. The base of the sapropel contains
an abrupt decrease in benthic fauna that continues through all of layer c. It also has an increase of the
coccolithophorids species Coccolithus pelagicus. Planktic foraminifera show at the same depth a peak of the cold
species Globorotalia scitula. These changes are followed by decreases in the carbonate preservation index and in
abundances of Globigerinita glutinata, Globigerinita uvula and Neogloboquadrina pachyderma (sinistral), which suggest
cold and highly productive upwelling waters. A short interval in the middle of the sapropel is characterized by low
values of C. pelagicus, a fluctuating increase of Pseudoemiliania lacunosa and among the foraminifera an increase of
Globigerinoides ruber together with the presence (although decreased) of G. glutinata, G. uvula and N. pachyderma
(sinistral). We interpret these features as suggesting high seasonality with warm stratified and probably oligotrophic
waters during summer and relatively cold conditions during winter. Finally, the topmost interval of the Vrica layer c
exhibits the re-appearance of P. lacunosa together with abundant siliceous phytoplankton. Planktic microfauna show
the disappearance of the cold species G. glutinata, G. uvula and N. pachyderma (sinistral). Thus this interval appears to
be characterized by warmer temperature. The transition from the laminated to the massive sediment displays a
sequence of events, including a decrease of the carbonate preservation index and peaks of Globorotalia inflata and
G. scitula, suggesting again upwelling and mixing of the whole water column and, thus, transition to the oxygenated
conditions characterizing the massive layer. Neither increased productivity nor stratification appear to characterize
the whole sapropel interval, which is, however, always dysoxic.
9 2002 Elsevier Science B.V. All rights reserved.

Keywords: calcareous nannofossils; foraminifera; paleoproductivity; Mediterranean; sapropel; Pliocene^Pleistocene

1. Introduction
* Corresponding author.
E-mail address: anegri@unian.it (A. Negri). The Vrica section is located 4 km south of the

0031-0182 / 02 / $ ^ see front matter 9 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 1 - 0 1 8 2 ( 0 2 ) 0 0 6 0 8 - 9

PALAEO 2969 24-12-02

244 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255
town of Crotone in southern Italy. The stratigra-
phy of this section is described by Bertolani Mar-
chetti et al. (1978), Colalongo et al. (1980), Na-
kagawa (1981), and Pasini and Colalongo (1994).
In 1985 the International Commission on Stratig-
raphy proposed this section as the stratotype for
the Plio^Pleistocene Boundary (Bassett, 1985).
However, we focused our attention on this section
because of its special paleoenvironmental features
and not its biostratigraphic signi¢cance. Sedimen-
tation in the Crotonese Basin occurred in a mod-
erately deep and restricted nearshore basin in
which the inferred water paleo-depth was between
500 and 800 m (Pasini and Colalongo, 1994). Sed-
imentation rates and thickness of the sedimentary
sequence are consequently magni¢ed. Sapropel in-
tervals here reach thicknesses of a meter or more ;
the c level in particular spans 3.38 m. In addition,
the sapropel intervals show a clear sub-milli-
metric lamination re£ecting annual sedimentation
(M. Claps, personal communication, 2001).
Decades after the ¢rst descriptions of sapropels
(Kullenberg, 1952; Olausson, 1961) and after doz-
ens of papers dealing with them, their origin is
still unsolved and under debate (see for instance,
Emeis et al., 1996; Rohling, 1999). Nonetheless,
there is general agreement that sapropel deposi-
tion is accompanied by conditions of anoxia or
dysoxia (Emeis et al., 1996; Cramp and O’Sulli-
van, 1999) and that the recurrent occurrence of
these layers is related to the Milankovitch preces-
sional cycle (Hilgen, 1991). However, some au-
thors consider enhanced primary production in
the photic zone to be primary among the factors
controlling concentration and accumulation of or-
ganic matter in the sea£oor and hence the major
cause of sapropel deposition (Calvert, 1983; Pe-
dersen and Calvert, 1990 among others).
Rohling (1994, 2001) argues that the formation
of eastern Mediterranean sapropels occurred in
conditions of anti-estuarine circulation that was
weakened relative to the present because of reduc-
tion of the eastern Mediterranean excess of evap-
oration over freshwater in£ux. This could lead to
a reduction of surface water salinities in the east-
ern Mediterranean, causing isolation of previously
formed high salinity (cooler) deep water. In this
way, mixing can be reduced and the consequently
PALAEO 2969 24-12-02
diminished oxygen advection could favor preser-
vation of organic matter. Further modeling by
Myers and Rohling (2000) supports this hypoth-
esis. Nevertheless, debate remains about what
triggers the deposition of sapropels.
The Vrica section represents a special opportu-
nity to evaluate in great detail the evidence for
those phenomena postulated to lead to sapropel
formation. Thanks to sub-millimetric lamination
that is present in the sapropels, the section may
provide the eventual possibility to reconstruct the
paleoceanographic history at an annual scale.
This paper presents new high-resolution micropa-
leontological data that contribute to unraveling
the sapropel problem.
2. Materials and methods
The Vrica core was drilled close to the classical
locality (Fig. 1) in March 1999 with a CMK MK
600 coring device that penetrated 50 m of sedi-
ment. The drilled succession shows a cluster of
sapropels corresponding to Vrica layers b, c, d,
and e (Selli et al., 1977; Pasini and Colalongo,
1994). After a careful sedimentological description
we focused on the thicker layer c as being ideal
for our purposes. This layer corresponds to inso-
lation cycle 180 that, according to Lourens et al.
(1996), has an orbitally tuned age of 1.851 Ma.
The laminated interval spans from 34.70 to 38.08
m. It shows laminae couplets having a mean
thickness of 0.5 mm that Claps (personal commu-
nication, 2001) interprets to represent annual de-
position (Fig. 2). These annual couplets imply a
time span of 6760 years for laminated layer c. The
section of the core containing layer c was sampled
every 5 cm in the laminated interval and every
10 cm in the massive intervals occurring above
and below the laminated interval. For calcareous
nannofossil analyses a smear slide was mounted
with Norland optical adhesive. No centrifuga-
tion was applied to concentrate the biogenic
fraction in order to retain the original com-
position of the nannofossil assemblage. Quanti-
tative analyses were performed with a light micro-
scope at 1250U magni¢cation on 126 samples
by counting at least 500 specimens/sample. The
 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255 245

Adriatic Sea

Tyrrhenian Sea VRICA

Ionian Sea

Fig. 1. Location map of the Vrica drill core.

abundance of nannofossils was converted to
number/mm2 , whereas the single species frequen-
cy was expressed in percent of total nannofos-
sils.
Samples for planktic and benthic foraminifera
analyses were dried at 60‡C, washed and sieved
with a 63 Wm mesh. The s 63 Wm fraction ob-
tained was examined for its planktic and benthic
foraminifera content. Where possible at least 300
specimens were counted in each sample, although
the generally low concentration of benthic forami-
nifera, especially in the sapropel samples, often
did not allow this. The relative frequencies of in-
dividual species are calculated as the percentage
of total planktic and benthic foraminifera fauna.
The concentration of planktic and benthic fora-
minifera (allochthonous and autochthonous
grouped separately) is calculated as number of
specimens per gram of dry sediment.
Following Howard and Prell (1994, and refer-
ences therein) we employed as a carbonate pres-
ervation index the (number of whole foraminif-
era)/(number of whole foraminifera + number of
fragments) U 100. This index remains the most
widely used and least ambiguous indicator of car-
bonate dissolution.

PALAEO 2969 24-12-02

246 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255

Fig. 2. Details of the laminated sediment characterizing the Vrica c-layer.

3. Results tive species showing interesting frequency £uctu-

3.1. Calcareous nannofossils
The calcareous nannofossil assemblage is well
preserved and abundant in both the massive
and sapropel layers. It consists mostly of abun-
dant Reticulofenestra spp. and ‘small’ Gephyro-
capsa spp. Other species like Pseudoemiliania la-
cunosa, Helicosphaera carteri, Helicosphaera sellii,
Syracosphaera pulchra, Rhabdosphaera claviger,
Rhabdosphaera stylifer, Coccolithus pelagicus, Cal-
cidiscus macintyrei, Calcidiscus leptoporus, Ponto-
sphaera spp., Holococcolithus spp. are less com-
mon. In general we noticed abundant reworked
specimens ranging in age from Late Cretaceous
to Miocene. In Fig. 3 we report the most indica-
ations within the sapropel. The most evident
feature is the decrease of the calcareous nanno-
fossil abundance in the interval from 34.36 to
38.23 m.
Another interesting feature is the C. pelagicus
abrupt increase between 38.23 and 38.08 m, where
after having shown very low values it reaches the
highest frequency. After this point it gradually
decreases to values that £uctuate but are not high-
er than 3.5%. P. lacunosa also shows a very inter-
esting pattern; it changes from values close to
10% below 38.08 m to values no higher than 2%
between 38.08 and 36.64 m. Above this point fre-
quency of the species strongly £uctuates until it
re-enters regularly in the assemblage at 35.78 m.
Other species like Helicosphaera carteri, Gephyro-

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 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255 247

Fig. 3. Relative abundances of selected calcareous nannofossil species across the c-layer (insolation cycle 180). On the left, note
the diagram showing the siliceous microfossil abundance. The di¡erent gray stripes re£ect the three phases evidenced in the lami-
nated interval.

capsa spp. and Reticulofenestra spp. show short-
term £uctuations in the laminated interval.
Finally, another feature observed while looking
through the smear slides is the abrupt occurrence
of siliceous fragments (centric and pennate dia-
toms, silico£agellates) starting from 36.68 m
(Fig. 4). The occurrence of those fragments con-
tinues until 34.70 m, and then they disappear.
3.2. Planktic foraminifera
In total 23 species and morphotypes were iden-
ti¢ed and used for interpretation. Counted cate-
gories are the species Globigerina bulloides, Globi-
gerina falconensis, Globigerina spp., Globigerinita
glutinata, Globigerinita uvula, Globigerinoides elon-
gatus, Globigerinoides gomitulus, Globigerinoides
ruber, Globigerinoides sacculifer, Globigerinoides
obliquus, Globoturborotalita apertura, Globoturbo-
rotalita rubescens, Globorotalia in£ata, Globorota-
lia scitula, Neogloboquadrina acostaensis, Neoglo-
boquadrina pachyderma, Orbulina universa, Turbo-
rotalita quinqueloba juveniles, planktic foraminif-
eral fragments, and teratological forms. Dextral
and sinistral forms of the Neogloboquadrinids
are counted separately. G. ruber, G. gomitulus
and G. elongatus are grouped together as G. ex
gr. ruber. The abundance pattern of the seven
most important categories is plotted in Fig. 4.
The carbonate preservation index indicates in-
creasing dissolution at the bottom and the top of
the laminated layer (37.93^37.63 m and 35.02^
34.70 m). Carbonate preservation is high in the
central part of the laminated sediment and in
the massive layers. G. uvula, N. pachyderma (sinis-
tral) and G. glutinata share the same trend. In
fact, they are present in the massive layer above
and below the laminated layer, but their abun-
dance quickly decreases between 38.08 and
37.71 m. After this point there is an evident in-
crease in abundance until 36.00 m. These species
almost completely disappear in the upper part of
the laminated layer, until the massive layer where
they re-occur with a percentage around 13%.
G. in£ata is present only in the massive intervals,
as well as G. scitula. However, the former species
disappears about 0.20 m (about 400 years) before
the deposition of the laminated layer, whereas

PALAEO 2969 24-12-02

248 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255

Fig. 4. Relative abundances of selected planktic and benthic foraminifera species across the c-layer (insolation cycle 180). The dif-
ferent gray stripes re£ect the three phases evidenced in the laminated interval.

G. scitula shows a strong peak at 38.04 m. G. ex
gr. ruber is usually low, with the exception of the
interval between 36.49 and 36.90 m where this
species abundance rises to values of about 12%.
G. bulloides is present throughout the core with
percentages lower than 5% of the total assem-
blage. Only two intervals (38.00^37.67 m and
35.60^34.60 m) record an increase in abundance
of this species to about 10% of the total assem-
blage.
3.3. Benthic foraminifera
In total 94 genera and species were identi¢ed.
Shallow-water epiphytic and epifaunal taxa such
as Elphidium spp., Asterigerinata spp., Discorbis
spp., Cibicides lobatulus, Buccella spp., Hanzawaia
boueana, Neocorbina spp., Planorbulina mediterra-
nensis, Rosalina bradyii, and Ammonia spp., are
grouped together as allochthonous, because these
species live exclusively in shallow water (Murray,
1991; Jorissen et al., 1993). We subtract the al-
lochtonous group from the total number of
benthic foraminifera, and subsequently we calcu-
late the number of benthic foraminifera per gram
of dry sediment (NBF). On the basis of ecological
requirements we selected some species that are
known not to tolerate low oxygen concentration.
This group consists of Cibicidoides kullenbergi,
Hoeglundina elengans, Oridorsalis spp. and Miliol-
ids.
We present in Fig. 4 plots of the NBF, the
group having high oxygen requirements and the
Epistominella exigua abundance. The NBF shows
a strongly decreasing abundance in the laminated
layer between 37.93 and 34.56 m where the mean
abundance is 15 specimens per gram dry sedi-
ment. In comparison, the NBF in the massive
layer is more than 100 specimens. The abundance
of the oxygenated taxa varies from 0 to 30%, but
it is evident that the higher values correspond to
the massive layer. In the laminated interval the
abundance of oxygenated taxa decreases and £uc-
tuates highly. Finally, E. exigua is almost absent
in the massive layer (38.80^37.93 m and between
34.59 m and the top). From 37.93 to 35.70 m its
abundance increases. We can recognize two inter-
vals, the ¢rst from 37.93 to 35.70 m that records
an increase of E. exigua reaching 20% of the total
abundance, and a second interval (35.70^34.59 m)

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 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255
in which relative abundance decreases to about
10%.
4. Discussion
The most signi¢cant micropaleontological fea-
ture observed in the laminated interval is the
abrupt decrease in the abundance of benthic fo-
raminifera across layer c. Comparison of the
benthic species composition in the massive layer
with recent and fossil data indicates a water pa-
leo-depth of about 700 m (Cita and Zocchi, 1978;
Murray, 1991). As stated in Section 3.3 we ¢nd
shallow-water species displaced from the near-
shore environment in the laminated sediments.
Shallow-water species are commonly found dis-
placed to greater depths in western Mediterranean
sediments (Jorissen, 1987). They are presumably
transported to deeper sites during storms. For this
reason we did not consider them autochthonous,
and therefore we subtracted this group from the
total number of benthic foraminifera. The lami-
nation, the absence of bioturbation, and the
strong decrease of NBF suggest stressed environ-
mental conditions and in particular very low oxy-
genation at the sea bottom.
The most striking feature observed in the cal-
careous nannofossil assemblage is its abrupt de-
crease across the laminated interval. This feature
was previously observed in sapropels by Negri et
al. (1999a,b), Negri and Villa (2000), and Negri
and Giunta (2001). Coccolitophorids generally
bloom in conditions of oligotrophy (Ziveri et al.,
1995a,b) and negatively correlate with diatoms
and foraminifera £uxes. Diatoms are actually
known as having a competitive advantage when
nutrient levels and supply rates are high (Ziveri et
al., 1995a,b; Sancetta, 1996). Sancetta (1994) pro-
posed that sapropels originated after enhanced
productivity caused by increased continental run-
o¡ in two ways: (1) by supplying high concentra-
tion of dissolved nutrients, and (2) because of
low salinity stratifying the surface waters. These
processes result in a bloom dominated by dia-
toms that under such conditions appear to be
the most competitive phytoplankton group. The
bloom occurs too rapidly for most zooplankton
PALAEO 2969 24-12-02
249
to respond and is thus not limited by grazing
pressure. Repetitions of diatom blooms on a sea-
sonal cycle over many years (Tang and Stott,
1993) could lead to the formation of packets of
laminae and the distinctive sapropel strata. On
this base Negri et al. (1999a) interpreted the cal-
careous nannofossil decrease characterizing sapro-
pels as a result of competition with more e⁄cient
£ora, such as diatoms.
In addition, the massive sediment di¡ers mark-
edly from the laminated interval by the presence
of the abundant planktic foraminifera G. in£ata
and G. scitula. G. in£ata is a transitional species
(Hemleben et al., 1989; Boltovskoy et al., 1996)
that dominates in well mixed waters having high
nutrient levels (Pujol and Vergnaud Grazzini,
1995). G. scitula is a polar^subpolar species (Be¤,
1977). Its presence is quite common in glacial-age
sediments of the Mediterranean Sea (Thunnel,
1978; Rohling et al., 1997; Capotondi et al.,
1998), although it occurs rarely in modern settings
(Pujol and Vergnaud-Grazzini, 1995). We can use
this species as an indicator of cold conditions.
Furthermore, its habitat depth ranges between
the surface and 1000 m (Be¤, 1980; Schiebel and
Hemleben, 2000). Itou et al. (2001) observed in a
North Paci¢c sediment trap that the abundance of
G. scitula does not necessarily correspond to sur-
face productivity and to particulate organic car-
bon (POC) £ux. Instead, it correlates well with
supply of ¢ne organic matter, which appears to
be a result of water convection. Thus, these au-
thors argue that the abundance of G. scitula
would basically correspond to intensi¢ed vertical
mixing. Based on these observations and on the
coeval increase of G. in£ata recorded in the Vrica
layer c, we interpret increased abundances of G.
scitula to re£ect the occurrence of a deep mixed
layer. Thus the main features di¡erentiating the
massive and the laminated intervals are the degree
of water column mixing and the availability of
dissolved oxygen at the sea bottom.
Details in the laminated interval allow descrip-
tion of the three subintervals that are summarized
in Fig. 5. At the base of the sapropel, changes
occur simultaneously in coccolith and foraminif-
era assemblages. Among coccoliths, we observe
the abrupt decrease of P. lacunosa and a sharp
250 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255

34.00 G. inflata disappears

34.26 G. scitula re-occurrence DEEP AND OXYGENATED MIXED LAYER
WELL OXYGENATED SEA BOTTOM
34.56 Repopulation benthic forams

34.70 G. inflata re-occurrence,

CPI increases, Diatoms disappear UPWELLING, CO2 RICH WATER

34.84 CPI decreases

35.78 reenter P. lacunosa SLIGHT REOXYGENATION

35.82 NBF slight increase OF THE SEA-BOTTOM

G. glutinata, G. uvula and WARM SUPERFICIAL WATER

35.92
N. pachyderma sin disappear
36.01 Drop G. uvula
36.12 Drop G. glutinata

36.35 G. ruber decreases

36.68 Diatoms occurrence

36.83 G. ruber and P. lacunosa increases STRATIFIED SUPERFICIAL WATER
HIGH SEASONALITY
36.86 C. pelagicus decreases
36.90 N. pachyderma sin decreases

G. glutinata, G. uvula and COLD SUPERFICIAL WATER

37.46
N. pachyderma sin occurrence STRONGLY DYSOXIC SEA BOTTOM

about
300m

37.63 CPI increases

UPWELLING IN THE
NBF strongly decrease, UPPER 300-400 M
37.93
CPI decreases INCREASING DISSOLUTION
38.00 G. scitula disappears
38.04 C. pelagicus and G. scitula peak INTERMEDIATE CONDITION
38.08 P. lacunosa disappears SLIGHT TURBULENCE, COLD
38.37 G. inflata disappears, G. scitula occurs
DEEP MIXED LAYER
C AT

38.52 G. inflata occurrence

Planktic foraminifera Benthic Foraminifera CPI = Carbonate Preservation

Index
Nannofossils Diatoms NBF = Number of Benthic
Foraminifera per dry gram sediment

Fig. 5. Reconstruction of the succession of main biotic events across the laminated c-layer of the Vrica drill core.

PALAEO 2969 24-12-02

 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255
peak of C. pelagicus. The latter coincides with a
sharp peak and the following abrupt decrease of
the deep-dwelling planktic foraminifera G. scitula.
C. pelagicus is a species known to live at high
latitudes in the northern hemisphere (Winter et
al., 1994), and thus has always been interpreted
as indicating low temperature (Ra⁄ and Rio,
1981). Recently, Cachao and Moita (2000) ob-
served C. pelagicus increasing in abundance close
to river mouths o¡ Portugal. This led the authors
to suggest that C. pelagicus abundance is related
to conditions of moderately high water turbulence
and nutrient availability.
These features suggest that the massive sedi-
ment/sapropel boundary records onset of cold
and mixed water. The sharp decrease of P. lacu-
nosa thus appears related to the occurrence of
cold water conditions at the beginning of sapropel
deposition. Unfortunately, this species became ex-
tinct in the late Pleistocene (254 kyr, Berggren et
al., 1995), and thus no data about its exact eco-
logical preferences are available. The decrease of
the carbonate preservation index observed in the
following interval (Fig. 4) corroborates the cold-
water hypothesis. As stated above, foraminiferal
fragmentation is one of the most widely used in-
dicators of carbonate dissolution (Thunnel, 1978;
Howard and Prell, 1994). This suggests the occur-
rence of highly corrosive waters, rich in CO2 , ris-
ing to the surface, probably as a consequence of
upwelling currents. Also, the peak of G. bulloides
supports this explanation. In fact, G. bulloides oc-
curs from the transitional to the polar zone (Be¤,
1977; Hemleben et al., 1989), and it lives in eu-
trophic areas when the surface mixed layer is rel-
atively deep (Thunnell and Reynolds, 1984; Kin-
caid et al., 2000). The almost complete absence of
benthic microfauna, especially in the ¢rst half of
the layer c, appears to con£ict with this hypoth-
esis. However, this apparent contradiction can be
explained by the fact that upwelling currents in
the Ionian Sea are known to be not deeper than
300^400 m (Russo, personal communication,
2002). Thus we can explain the coexistence of
two water masses characterizing di¡erent environ-
ments : a deeper one ( s 400 m depth), in which
we observe strong dysoxia, and a shallower one
that sees the rising of CO2 -rich intermediate cold
PALAEO 2969 24-12-02
251
water, probably originating in the Northern Adri-
atic (Adriatic Intermediate Water, AIW) as sug-
gested by Rohling (1994, 2001). Another impor-
tant feature observed in the whole sapropel with
respect to the massive intervals is the relative in-
crease of E. exigua among the scarce benthic
assemblage. This species is reported in dysoxic
environments (Jorissen et al., 1993), but it is
also common in North Atlantic locations where
this species quickly responds to phytoplankton
blooms (Gooday, 1988). Thus we interpret this
feature as supporting the hypothesis of an in-
crease in primary productivity. The increase of
nitrogen ¢xation associated with layer c as postu-
lated by Arnaboldi and Meyers (in press) sup-
ports this hypothesis.
At 37.46 m, we detected the appearance of the
species G. glutinata, G. uvula and N. pachyderma
(sinistral). Schiebel and Hemleben (2000) ob-
served that the dominance of G. glutinata in the
North Atlantic (about 50‡N) coincides with a
deepening of the mixed layer and hence with a
decreased surface water temperature. Thus G. glu-
tinata can be considered a species related to cold
conditions. Although observations on G. uvula in
modern assemblage are rare (Boltovskoy et al.,
1996; Mu«nke and Hemleben, 1999), this species
is generally found in cold waters (Boltovskoy et
al., 1996). N. pachyderma (sinistral) is known to
be the only species living at high latitudes (Hem-
leben et al., 1989) and it tolerates temperatures
not higher than 15‡C. Thus it appear that the
assemblage consisting of N. pachyderma (sinistral)
and G. glutinata supports the hypothesis of low
water temperature characterizing the ¢rst part of
the layer c depositional history. The co-occur-
rence of G. uvula is probably related to the same
conditions.
The second subinterval spans approximately
1 m (from 36.90 to 35.92 m, Fig. 5). It is charac-
terized by an increase in frequency of the planktic
foraminifera G. ruber together with the continued
presence, although in lower abundances, of G.
glutinata, G. uvula and N. pachyderma (sinistral).
G. ruber is a symbiont-bearing species, living in
tropical^subtropical areas, that tolerates temper-
atures between 14 and 32‡C (Bijma et al., 1990).
According to Kincaid et al. (2000), it is present in
252 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255
the Santa Barbara Basin only when surface waters
are warmest (from 14 to 19.8‡C), with £uxes in-
creasing when sea-surface temperatures increase.
Increased £uxes co-occurred with increased sili-
ceous plankton contributions from warm oceanic
diatoms and warm-water radiolarians. It is note-
worthy that our analyses identi¢ed the abrupt ap-
pearance in the c-layer of diatoms and siliceous
fragments coinciding with the beginning of the G.
ruber increase. However, we can also invoke high
seasonality in order to explain the co-occurrence
of the cold species N. pachyderma (sinistral) and
G. uvula, which probably contribute to the assem-
blage during cold winters. Also, this interval re-
cords the relatively high abundance of E. exigua,
testifying that productivity was high.
Finally, the third subinterval, spanning 1.22 m
(Fig. 5), records the almost complete disappear-
ance of the cold planktic foraminifera N. pachy-
derma (sinistral), G. uvula and G. glutinata. These
signals suggest warmer sea-surface temperatures.
The phytoplankton community records a slight
decrease of Helicosphaera spp. and the continuous
and increasing presence of P. lacunosa, diatoms
and in general of the siliceous fragments. In mod-
ern coccolitophorid assemblages Helicosphaera
spp. behavior is not clearly understood. In fact,
Pujos (1992) suggested that increased frequencies
of H. carteri indicate a sudden decrease in salinity
and/or an increase in productivity. Furthermore,
Haidar and Thierstein (1997) recorded in modern
Bermuda coccolitophore assemblages that H. car-
teri increases under conditions of high tempera-
ture and light intensity and when nutrients are
lowered with respect to the amount required by
Emiliania huxleyi, and Ziveri et al. (1995a,b) ob-
served increased £uxes of H. carteri when overall
coccolithophore £uxes were high.
P. lacunosa became extinct during the Late
Pleistocene, but it is a placolith-bearing species.
In modern assemblages, placolith-bearing species
dominate in coastal and upwelling areas (Young,
1994). Therefore it appears that the calcareous
nannofossils suggest enhanced productivity (in-
crease of P. lacunosa). The diatom increase (Fig.
4) appears to further support this hypothesis.
The decreasing carbonate preservation index at
the top of the sapropel (34.84^34.70 m) again
PALAEO 2969 24-12-02
suggests the rising of deep, cold, CO2 -rich waters,
probably re£ecting upwelling currents. Also, in
this case the slight increase of G. bulloides sup-
ports this hypothesis. This event precedes the res-
toration of the normal conditions characterizing
the massive interval above the layer c. It probably
represents the trigger that led to water mixing into
the deeper environments. This resulted in the re-
occurrence of the deep-dweller forms (G. scitula
and G. in£ata), the disappearance of the siliceous
phytoplankton, and sea bottom repopulation by
benthic foraminifera.
To summarize, Vrica sapropel layer c formed
during three di¡erent phases evident in the micro-
paleontological assemblages. It is important to
point out the almost complete independence of
super¢cial and bottom water and in particular
that strati¢cation (de¢ned as the presence of a
fresh-water super¢cial layer inhibiting the mixing
at greater depth sensu Vergnaud-Grazzini et al.,
1977) does not occur at the beginning of sapropel
deposition. Strati¢cation is probably stronger
only in the middle part of layer c. An increase
in productivity characterizes the whole sapropel,
but at the beginning it is not stronger. Thus a
model of sapropel deposition based exclusively
on increased productivity (Pedersen and Calvert,
1990) appears not justi¢ed.
A sort of strati¢cation occurs in the sense that
two di¡erent water-masses coexist. However,
whereas super¢cial waters document upwelling
alternating with strati¢cation, low oxygenation
conditions at the sea bottom persist, probably
because of the cessation of ADW formation in-
voked by Rohling (1994, 2001). In conclusion, our
results point to dysoxia at the sea bottom as the
main reason for sapropel formation. This mecha-
nism is probably the same as for younger sapro-
pels.
5. Conclusion
Our study identi¢es three di¡erent paleoceano-
graphic phases recorded in the Vrica c-layer,
which corresponds to insolation cycle 180 (1.851
Ma). In general, our data show that high produc-
tivity and enhanced water strati¢cation did not
 A. Negri et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 190 (2003) 243^255
coexist during sapropel deposition. The ¢rst phase
is characterized by the occurrence of strong dys-
oxia at the sea bottom. This was probably caused
by diminished formation of cold and saline waters
in the North Adriatic Sea and slowing of their
subsequent southward £ow (Rohling, 1994,
2001). This cessation led to the almost complete
oxygen depletion at the sea bottom in the Crotone
Basin of the Ionian Sea. However, in this area, at
the same time, upwelling a¡ected the upper 300^
400 m of the water column and caused slightly
enhanced productivity (mesotrophic conditions,
but not eutrophic).
The second phase exhibits the high productivity
and strong water strati¢cation that are typical
features of other sapropels, together with strong
seasonality. Finally, the third phase records
warmer sea-surface temperatures and probably
higher productivity. The transition to the oxygen-
ated massive interval appears to pass through an
upwelling event a¡ecting only the super¢cial
water column. Oxygenated conditions at the sea
bottom are probably related to restoration of the
North Adriatic Deep Water formation.
We postulate that two mechanisms are needed
to explain the formation of the c-layer. The ¢rst is
at the basinal level and involves diminished for-
mation of the North Adriatic Deep Water £ow, in
agreement with Rohling (1994, 2001). The slowed
circulation leads to oxygen depletion and dysoxia
or anoxia at the sea bottom. It is probably the
mechanism responsible for the formation of all
the sapropels in the Mediterranean Sea. A second
mechanism ^ upwelling currents ^ appears related
to local wind stress and the physiography of the
Ionian Basin. It probably triggers increased pro-
ductivity, but anoxia (and thus increased preser-
vation) seems to exist from the beginning of the
sapropel. Water strati¢cation occurs after the
laminated interval has already started to deposit.
Thus, high productivity appears not to be the
trigger for sapropel deposition.
Acknowledgements
We are indebted to Massimo Sarti for conceiv-
ing and organizing drilling of a core at the Vrica
PALAEO 2969 24-12-02
253
Site and Michele Claps for help in core logging.
Comments of Phil Meyers, Mike Howell and an
anonymous reviewer helped to improve our man-
uscript.
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