ELSEVIER Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156

Middle to Upper Miocene benthonic foraminiferal palaeoecology of

the Tap Marls (Alicante Province, SE Spain) and its
palaeoceanographic implications
Holger Gebhardt *
Department of Geology and Mineral Sciences, University of Ilorin, P.M.B. 1515, Ilorin, Kwara State, Nigeria
Received 30 December 1997; accepted 3 June 1998

Abstract

The benthonic foraminiferal faunas of twenty sections have been analysed statistically (cluster analysis) and interpreted
palaeoecologically. The analysis shows that oxygenation was the most important factor for the distribution of the faunas in
the investigated area. The lifting and lowering of the sill to the Atlantic Ocean (Gibraltar sill), in conjunction with climatic
changes and sea level fluctuations, led to changes in the current system and to two faunal turnovers at the transition from
Langhian to Serravallian (15.215.0 Ma) and at the end of the Serravallian, respectively (11.711.4 Ma). The faunas
indicate well oxygenated (Langhian), reduced oxygenated (Serravallian) and again well oxygenated (Tortonian) conditions.
A simplified circulation model for the western Mediterranean, based on the authors interpretation and additional sources,
suggests: (a) an estuarine circulation during the Langhian; (b) a restricted anti-estuarine pattern with sluggish circulation
until the end of the Serravallian; and (c) an anti-estuarine circulation similar to todays situation during the Tortonian.
1999 Elsevier Science B.V. All rights reserved.

Keywords: Middle to Upper Miocene; benthonic foraminifera; palaeoecology; southeastern Spain; palaeoceanography;
circulation patterns; western Mediterranean

1. Introduction or seasonality (e.g. Katz and Thunell, 1984; Van

Benthonic foraminiferal distribution may be re-
lated to water masses on a large scale (e.g. Lohmann,
1978; Douglas and Woodruff, 1981; Culver, 1988;
Denne and Sen Gupta, 1991), but the occupation of
different microhabitats is governed by a more local
(or regional) interplay of nutrient (food) availabil-
ity, oxygen supply (both are frequently combined)
Present address: Technische Universitat Berlin, Institut fur
Angewandte Geowissenschaften II, Sekr. EB 10, Ernst-Reuter-
Platz 1, D-10587 Berlin, Germany.
der Zwaan and Jorissen, 1991). Other factors like
water depth or salinity are also reflected in the fo-
raminiferal record and have to be considered in the
interpretation. Benthonic foraminifera represent one
of the most sensitive indicators of dissolved oxygen
levels (Van der Zwaan and Jorissen, 1991; Kaiho,
1994) and can therefore be used to interpret ancient
sediments. However, fossil assemblages are more
difficult to interpret in terms of water-mass charac-
teristics as small-scale differences in the physical and
chemical particularities of overlying waters may not
affect the fossil record distinctly. Bioturbation and

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142 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156

Fig. 1. Location of the investigated sections (AT), single samples (E4; Tp1,2). Outcrops of Langhian to Tortonian rocks modified
after Amela Samper et al. (1975), Martnez et al. (1975, 1978), Colodron et al. (1981) and several unpublished M.Sc. theses from the
Technische Universitat Berlin (19791992).

sampling technique may also narrow the resolution
power to such an extent that shorter environmental
changes may not become obvious. This study, among
others, makes a first step to create a general palaeo-
ceanographic model of the western Mediterranean
for the Langhian to pre-Messinian time interval.
Gebhardt (1994) did extensive investigations on
the foraminiferal faunas of the Tap Marls of the
Alicante Province in southeastern Spain. Considera-
tion of additional and recently published literature,
in particular about stable carbon and oxygen isotope
data and the application of a newly developed oxy-
gen indicator method (Kaiho, 1994), makes a new
interpretation of his benthonic foraminiferal counts
necessary. An attempt is made to fit the new results
into a palaeoceanographic model.
The investigated area covers large parts of the
geological maps of Castalla (Martnez et al., 1978),
Onteniente (Martnez et al., 1975), Alcoy (Amela
Samper et al., 1975) and of Villajoyosa (Colodron
et al., 1981), all situated in the northern part of the
Alicante Province. The localities of the sections and
single samples are given in Fig. 1.
The biostratigraphic ages of the investigated sec-
tions have been determined by means of planktonic
foraminifera, using the zonation of Iaccarino and Sal-
vatorini (1982), which provides the highest strati-
graphic resolution in the Mediterranean. This zona-
tion has been correlated with a recently published
timescale (Berggren et al., 1995). The earliest evi-
dence of Tap Marl sedimentation has been recorded as
uppermost Burdigalian, but major deposition started
in the basal Langhian and ended in the investigated
region within the Tortonian (see Gebhardt, 1994).
 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
2. Geological and tectonic setting of the study
area
Neogene whitish marls and clays are called Tap
facies in the eastern part of the Betic Cordillera,
corresponding to Albarizas or Moronitas facies of
the western parts. Over large parts of the Alicante
Province, the Tap Marls are divided by a turbiditic
or debris flow stack of calcareous conglomerates and
arenites (Cater, 1987; Gebhardt, 1994). In the above-
mentioned geological maps, this unit is referred to as
Transgression serravaliense. Therefore, a Langhian
Tap 1 can be distinguished from a Serravallian to
Tortonian Tap 2 (see Fig. 2). The turbidite series
represents the peak of tectonic activity (Fig. 2).
Generally, basin development is controlled by
folding and the diapirism of underlying salifer-
ous rocks (Sanz de Galdeano and Vera, 1992). Ott
dEstevou et al. (1988) have reported three changes
in the palaeostress field during the Miocene. The tec-
tonic activity has led to stronger deformation of the
Tap 1 and the turbidites, including submarine erosion
of the Tap 1 (Ott dEstevou et al., 1988; Gebhardt,
1994), whereas the Tap 2 deformation appears more
Fig. 2. Schematic lithologic section of investigated Miocene
rocks in the northern part of the Alicante Province.
143
gentle. The strike of fold axes is generally ENE
WSW, which is also visible from the basin arrange-
ment and outcrop of Langhian to Tortonian rocks
as shown in Fig. 1. Diapirism as well as abnormal
trending fold axes in the eastern part of the inves-
tigated region can be explained with a combination
of shortening and dextral wrenching, which also pro-
duced the pull-apart structures in which the diapirs
then could intrude (De Ruig, 1990).
The marls are dark grey to brownish in colour
if unweathered. They alter to blue, green, grey and
whitish colours when they have been exposed to
weathering. The marls are probably laminated, al-
though they may appear massive if they are fresh.
Bioturbation has been observed only in or between
nearby turbiditic calcarenite layers. No macrofossils
have been observed within the marl facies. However,
the turbiditic layers often contain large amounts of
shallow-water organic remains (e.g. calcareous al-
gae, bivalves, etc.). Additional faunal and floral com-
ponents of the marls are sponge spicules, ostracods,
radiolaria, fish teeth and a probably larger content of
coccoliths. The carbonate content of the marls varies
between 26% and 94%, average 60% (Gebhardt,
1994).
3. Material and methods
Gebhardt (1994) investigated foraminiferal faunas
of 190 marl samples from 20 sections and 3 single
samples (Fig. 1). All samples were dried, treated
with hydrogen peroxide and washed over a 0.100
mm sieve. After determination of the P=B ratio (con-
tent of planktonic foraminifera), the residues were
dry sieved over a 0.250 mm sieve. An amount of
about 300 specimen of benthonic foraminifera has
been picked out of the >0.250 mm fraction where
possible. The complete data set (absolute and per-
centage occurrences), including logs of the sections
can be obtained from Gebhardt (1994).
Although Schroder et al. (1987) have criticized
the restriction to larger fractions, mainly because of
the possible loss of potential environmental index
species, this restriction has a lot of advantages in
practice and allows a proper analysis and interpre-
tation of palaeoenvironments because faunal bound-
aries remain the same in both, the >0.063 mm and
 144
Table 1
Major benthonic foraminiferal associations and their most frequent species together with their assumed palaeoenvironments and further allocatable minor associations

Association: Cibicidoides pachyderma Melonis pompilioides= Melonis pompilioides= Globobulimina pyrula

H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156

Cibicidoides dutemplei Siphonina reticulata
No. of samples: 22 82 40 3
max.% max. % max. % max. %
Most frequent Cibicidoides pachyderma 45 Melonis pompilioides 45 Melonis pompilioides 39 Globobulimina pyrula 44
benthonic foraminifera Lenticulina stellata 26 Cibicidoides dutemplei 37 Siphonina reticulata 31 Praeglobobulimina
Melonis pompilioides 25 Cibicidoides pachyderma 29 Melonis barleeanus 28 affinis 17
Uvigerina barbatula 21 Cibicidoides subhaidingeri 19 Gyroidinoides peramplus 26 Chilostomella ovoidea 9
Uvigerina hispida 19 Uvigerina barbatula 18 Nodosaria longiscata 22
Siphonina reticulata 17 Praeglobobulimina pupoides 17 Islandiella laevigata 20
Cibicidoides Nonion boueanum 14 Stilostomella consobrina
pseudoungerianus 16 Siphonina reticulata 13 emaciata 17
Pullenia bulloides 15 Martinottiella communis 12 Haplophragmoides sp. 17
Saracenaria italica 11 Praeglobobulimina affinis 11 Cibicidoides pachyderma 15
Islandiella laevigata 9 Stilostomella monilis laevigata 14
Additional allocatable Uvigerina spp., 1 sample Valvulineria complanata, Siphonina reticulata, 3 samples
minor associations 1 sample Melonis barleeanus, 1 sample
Cibicidoides dutemplei, Globulina gibba, 1 sample
1 sample Ammobaculites humboldti latum,
Lenticulina spp., 9 samples 1 sample
Lenticulina spp., 3 samples
Average content of 3% dysoxic 6% dysoxic 3% dysoxic 68% dysoxic
oxygen indicators a 73% suboxic 57% suboxic 68% suboxic 20% suboxic
25% oxic 34% oxic 29% oxic 12% oxic
Assumed well oxygenated reduced oxygenation well oxygenated oxygen deficiency
palaeoenvironment (high oxic) (low oxic) (high oxic) (dysoxic)
12% middle bathyal 84% middle bathyal 63% middle bathyal middle bathyal
88% lower bathyal 16% lower bathyal 37% lower bathyal

The percentages for the bathyal zones correspond to the co-occurrence of association and depositional environment in years for each section.
a Only >0.250 mm pickings were evaluated. Therefore, percentages of oxic indicators (large) are over-emphasized as well as dysoxic indicators (usually small) are

underrepresented.
 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
the >0.250 mm fractions (Lutze, 1980). However,
it may restrict the direct comparability with other
studies. The method has been successfully applied
before, e.g. Lohmann (1978), Lutze and Coulbourn
(1984) or Caralp (1988). The major advantages are:
(1) reduced effects of transport by bottom currents,
(2) easier classification, and (3) significant reduction
of working time required. These are real advantages
in the Gebhardt (1994) study because of the some-
times high amounts of displaced and reworked spec-
imen from shallower waters or older deposits (up to
40% of the fauna). Recrystallization as well as partial
dissolution may hinder the comparison of sections in
this tectonically stressed area of investigation, as
smaller forms are sometimes not recognizable.
To create groups of samples of similar faunas,
Gebhardt (1994) applied a Q-mode cluster analy-
sis. The created clusters represent different thanato-
coenoses which in turn can be allocated to different
palaeoenvironments. The analysis was done with the
SPSS=PCC program in the average linkage-fusion
modus to avoid modus-related influences on the clus-
tering process as much as possible (see also Back-
haus et al., 1994; Shi, 1993). For the computation,
percentages of species, x, were transformed to ln.x/
to promote normalization; Squared Euclidean dis-
tance has been used as similarity index (standard
setting of SPSS=PCC). To maintain the representa-
tivity of the samples, minimum sample size was 80
individuals and only species with a minimum con-
tent of 5% in at least one sample have been chosen.
Of course, displaced and reworked foraminifera have
been excluded from the cluster analysis. By applying
the elbow criterion (Backhaus et al., 1994), a total
of eleven cluster have been created. They represent
four major associations of varying sizes and seven
minor associations of 13 samples, which have been
allocated to the major associations because of their
common palaeoenvironments (Table 1). Because of
dominance of non-indicative species, two clusters
contain faunas of different environmental conditions.
Their samples had to be sorted and allocated by
hand to two of the major associations. As a result
of this experience, cluster analysis is a good tool to
handle large data sets, but much attention has to be
paid in their interpretation. The additional applica-
tion of an environmental indicator species concept is
absolutely necessary.
145
4. Microhabitats
All species classified in the cluster analysis are
indicative of more or less normal marine salinities.
At least, salinity seems to have no influence on
the clustering process. Therefore, only oxygenation
and water depth as factors with influence on the
distribution of the benthonic foraminifera of the Tap
Marls will be discussed here.
Availability of food (Dorganic matter) and oxy-
gen are the most important parameters for fora-
minifera (Sjoerdsma and Van der Zwaan, 1992).
Both are usually closely related: the higher the or-
ganic matter supply, the lower the oxygen content
in the water. The decay of organic matter may lead
to the presence of extensive zones of anoxic or
dysoxic conditions at the sedimentwater interface.
The amount of organic supply, on which bathyal
benthonic foraminifera largely depend, in turn de-
pends on mineral nutrients (mainly phosphate and
nitrate) and surface productivity (for further discus-
sion see Brasier, 1995). Additional important sources
of organic matter or excess nutrient input are river
discharge or man-made sewage (e.g. Seiglie, 1968;
Jorissen, 1987; Van der Zwaan and Jorissen, 1991).
However, a large-scale direct impact of such sources
on the benthonic foraminiferal fauna is probably re-
stricted to wide shelf areas (Van der Zwaan and
Jorissen, 1991).
With increasing food supply, or decreasing pres-
ence of free oxygen, respectively, r-selected (op-
portunistic) species are favoured over K-selected
(equilibrium) species. The organic flux plays also a
dominating role in deciding the number of benthonic
foraminifera (high flux allows high standing stocks).
Its variation however seems to become a subordinate
factor below 500 m water depth (Sjoerdsma and Van
der Zwaan, 1992).
Since there is no general consensus among sci-
entists about subdivision of oxygen levels (Tyson
and Pearson, 1991 discussed this topic intensively),
the definition of Kaiho (1994) is used in this paper:
anoxic (00.1 ml=l O2 ), dysoxic (0.10.3 ml=l O2 ),
suboxic (0.31.5 ml=l O2 ), low oxic (1.53.0 ml=l
O2 ), and high oxic (>3.0 ml=l O2 ).
Foraminifera are oxygen consumers which can,
generally, tolerate significant variations in dissolved
oxygen in the ambient water. Benthonic foraminifera
146 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
utilize the oxygen at the sedimentwater interface
and of pore waters within the surface sediments. The
presence of sulphides is toxic to organisms other than
anaerobic bacteria, but some foraminifera are present
in the redox-potential-discontinuity layer, which con-
tains both O2 and H2 S. However, no facultative
anaerobes among foraminifera are known, but some
species may tolerate periodical anoxia (Sen Gupta
and Machain-Castillo, 1993). Such small, thin, unor-
namented, highly perforate test-bearing foraminifera
should survive short-lived anoxic periods whereas
others would not (Bernhard, 1986). Benthonic fo-
raminifera are absent under conditions without any
oxygen (Murray, 1991), at least if such conditions
remain constant for longer periods.
Corliss (1985) investigated the distribution of
bathyal to abyssal, living foraminifera in the sedi-
ments of box cores. He distinguished between those
species with epifaunal preference (Oridorsalis tener,
Fontbotia wuellerstorfi), which dominate the top 1
cm of the sediment (Dshallow infaunal, Buzas et
al., 1993, see below) and those with infaunal pref-
erence (Melonis barleeanum, Globobulimina affinis,
Chilostomella oolina). The latter two species live
from 6 to 15 cm depth, whereas Melonis barleeanum
dominates the 2 to 3 cm interval. He found that the
species distribution is not directly controlled by the
overlying bottom water but is influenced by varia-
tions of the pore-water oxygen content. Thus, deeper-
dwelling species will migrate closer to the surface if
the oxygen content of the bottom water decreases. In
a later study (Corliss, 1991), a subdivision into epi-
faunal (01 cm), shallow infaunal (02 cm), interme-
diate infaunal (14 cm) and deep infaunal (>4 cm)
was made. Buzas et al. (1993) evaluated a larger set
of data statistically and distinguished between shal-
low infaunal (maximum density in the top 2 cm of the
sediment) and deep infaunal (maximum density be-
low the top 2 cm of sediment). These groups (micro-
habitats) are represented by distinct morphological
characteristics which suggest its application to fos-
sil communities. Additionally, the deep infaunal taxa
are found at progressively shallower sediment depth
with increasing organic carbon content, which may
correlate with corresponding lower oxygen contents.
Arenaceous taxa are found to be mostly deep infaunal
(Buzas et al., 1993), probably indicating their greater
ability to cope with lower oxygen values.
The concept of correlation of fauna, sediment
depth and oxygen content was developed further
by Kaiho (1994). Although his paper can be criti-
cized because of its simplifications, it provides an
applicable concept for the interpretation of ancient
environments.
With decreasing oxygen content, deep infaunal
forms become shallow infaunal, or even epifaunal
with total anoxia in the sediment and little oxygen in
the bottom water (Kaiho, 1991, 1994; Van der Zwaan
and Jorissen, 1991). Kaiho (1994) distinguished be-
tween three categories of dissolved oxygen indica-
tors (only taxa found in the Tap Marls are mentioned
here).
(1) Dysoxic indicators, characterizing oxygen
contents of 0.1 to 0.3 ml=l. These are thin-walled,
elongate, flattened, mostly small species of Bolivina,
Bulimina, Cassidulina, Chilostomella, Dentalina,
Stilostomella, Pleurostomella, Globobulimina (Prae-
globobulimina), which also live as infauna under
highly oxygenated bottom waters. Their thin, highly
porous, small tests and the high surface=volume ra-
tios may provide for improved mitochondrial oxygen
uptake (Leutenegger and Hansen, 1979, in Sen Gupta
and Machain-Castillo, 1993) or indicate difficulties
in calcium carbonate secretion (Phleger and Soutar,
1973). This group corresponds in part to the deep
infaunal group of Corliss (1991); some taxa are also
found in shallow to intermediate infauna microhabi-
tats.
(2) Oxic indicators, pointing to oxygen contents
of >1.5 ml=l. These are generally larger than 0.350
mm, thick-walled epifauna with planoconvex, bicon-
vex, rounded trochospiral or spherical tests such as
Cibicidoides spp., Fontbotia wuellerstorfi, Siphonina
reticulata (see Van der Zwaan, 1982). The epifaunal
taxa of Corliss (1991) correspond to this group. Such
species are absent in low-oxygen environments.
(3) Suboxic indicators, characteristic for oxygen
contents of 0.3 to 1.5 ml=l. They include oxic indica-
tors smaller than 0.350 mm, certain species of Bulim-
ina, Cassidulina, Fissurina, Gyroidinoides, Lagena,
Lenticulina, Melonis, Nonion, Oridorsalis, Pullenia,
Sphaeroidina bulloides, Uvigerina, Valvulineria and
thick-walled Dentalina as well as thin-walled species
found in intermediate microhabitats close to dysoxic
indices. These taxa prefer shallow to intermediate
infaunal microhabitats (compare Corliss, 1991).
 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
As a consequence, oxygen deficiency in the bot-
tom water can be proved only by the absence of oxic
indicators (or with further depletion of oxygen by
absence of suboxic indicators) and not by the pres-
ence of species with tolerance to oxygen deficiency
(see also Kaiho, 1991). This would be another ar-
gument for the restriction to larger fractions of the
fauna (e.g. >0.250 mm) because the existence of
large specimens of certain taxa is only indicative
for a higher oxygen supply at the watersediment
interface. Inclusion of the smaller fractions results
in a shift towards the smaller taxa (in this study:
Bolivina, Bulimina, Trifarina and Uvigerina, all in-
faunal genera; Murray, 1991) which then represent
the conditions in the sediment rather than at the
watersediment interface.
5. Benthonic foraminiferal assemblages
The cluster analysis has merged the foraminiferal
faunas to eleven groups. The four largest represent
major assemblages, and the remainder are minor
assemblages of one to three samples which could
easily be allocated to the major associations due to
similar ecologic characteristics of their faunas. Ta-
ble 1 shows the major assemblages together with the
most frequent species. The most important ecologic
factor for the separation of the assemblages is oxy-
genation. Other factors such as salinity, water depth
or temperature are not expressed by this grouping.
At least, no significant hint can be derived from the
foraminiferal content of the assemblages for these
factors. Therefore, they can be neglected for the in-
terpretation of the clusters. Criteria to estimate oxy-
genation are based on foraminiferal morphology, test
size, wall thickness or indicative taxa (e.g. Phleger
and Soutar, 1973; Bernhard, 1986; Kaiho, 1994). In
addition to the foraminiferal fauna, some sedimen-
tological characteristics and the trace fossil content
are used to estimate the oxygen content of the an-
cient bottom water. Savrda et al. (1984), Savrda and
Bottjer (1986) and also Vercoutere et al. (1987) have
shown a positive correlation between burrow size
and oxygen content of bottom waters.
Stress-tolerant species with opportunistic life
strategies in this study, which can tolerate high or-
ganic flux or low-oxygen supply, belong to the gen-
147
era Bulimina, Chilostomella, Globobulimina, Prae-
globobulimina, Uvigerina and Valvulineria (Phleger
and Soutar, 1973; Douglas and Heitman, 1979;
Mullineaux and Lohmann, 1981; Van der Zwaan,
1982, 1983; Miller and Lohmann, 1982; Bernhard,
1986; Jorissen, 1987; Meulenkamp and Van der
Zwaan, 1990). The frequent occurrence of Cibici-
doides (in particular C. dutemplei and C. pseudoun-
gerianus) and Melonis is attributed to high food
supply by some authors (Arnold, 1983, in Caralp,
1988; Meulenkamp and Van der Zwaan, 1990).
The associations are named by their most frequent
species. In two cases, the second most frequent
species name was included to distinguish between
them. Only the frequent species are mentioned here.
Note that arenaceous species have not been included
in the calculations of percentages.
5.1. Melonis pompilioides=Siphonina reticulata
association
This assemblage is characterized by a large
number of oxic indicators such as Siphonina
reticulata or Cibicidoides pachyderma (457%,
average 29%) and of suboxic indicators (No-
dosaria longiscata, Melonis pompilioides, Melonis
barleeanus, Gyroidinoides peramplus, Islandiella
laevigata, Stilostomella consobrina var. emaciata,
Stilostomella monilis var. laevigata; 4398%, av-
erage 68%). Dysoxic indicators occur only occa-
sionally in the >0.250 mm fraction (mainly Prae-
globobulimina pupoides; 020%, average 3%). This
assemblage is restricted to the Tap 1 marls. Its sed-
iments are usually whitish or bright green and only
slightly compacted. Bioturbation is not visible and
weathered Tap 1 marls appear to be laminated. How-
ever, this appearance may be due to compaction
during diagenesis. Bioturbation can be observed in
some places inside the frequent turbiditic calcarenite
layers. Sometimes these layers are connected by cal-
carenite-filled burrows of relatively big size (several
cm in diameter). Some minor associations have been
allocated to this association because of its lack of
dysoxic indicators and strong dominance of oxic in-
dicators (Table 1). Thus, well oxygenated conditions
corresponding to the high oxic level of Kaiho (1994)
are indicated for the samples of this association.
148 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
5.2. Melonis pompilioides=Cibicidoides dutemplei
association
Also in this association, oxic indicators are fre-
quent (Cibicidoides dutemplei, Cibicidoides pachy-
derma, Cibicidoides subhaidingeri, Siphonina retic-
ulata; 469%, average 34%) as well as suboxic
indicators (Melonis pompilioides, Uvigerina barbat-
ula, Nonion boueanum; 3189%, average 57%).
However, this assemblage contains also significant
amounts of dysoxic indicators such as Praeglobob-
ulimina pupoides and Praeglobobulimina affinis (0
39%, average 6%). This association occurs mainly
in the Serravallian beds of Tap 2, which are charac-
terized by dark grey to bluish, strongly compacted
and frequently H2 S-smelling marls. Although the
marls look generally massive if fresh, lamination
occurs in some places. Visible bioturbation is re-
stricted to turbiditic layers and the burrows are dis-
tinctively smaller in size than those of the Melonis
pompilioides=Siphonina reticulata association. Some
minor associations have been allocated to this as-
sociation because of the relatively high amounts of
dysoxic indicators in this samples (Table 1). It should
also be considered that only >0.250 mm pickings
were evaluated. Therefore, oxic indicators (large)
are overemphasized as well as dysoxic indicators
(usually small) are under-represented in the count-
ings. Because percentage calculations are based on
the subdivisions of Kaiho (1994), Cibicidoides was
counted as oxic indicator. This genus, in particular C.
dutemplei, is assumed to tolerate little oxygen defi-
ciency or increased nutrient supply (Van der Zwaan,
1982; Meulenkamp and Van der Zwaan, 1990). The
Melonis pompilioides=Cibicidoides dutemplei asso-
ciation, may therefore contain less pure oxic in-
dicators than the percentage value suggests. Thus,
a reduced oxygen supply can be assumed for this
association. However, since oxygen indicators are
frequent, this association may represent the low oxic
level.
5.3. Cibicidoides pachyderma association
This assemblage contains almost only oxic indica-
tors (Cibicidoides pachyderma, Siphonina reticulata,
Cibicidoides pseudoungerianus; 1250%, average
25%) and suboxic indicators (Lenticulina stellata,
Melonis pompilioides, Uvigerina barbatula, Uvige-
rina hispida, Pullenia bulloides, Saracenaria ital-
ica, Islandiella laevigata; 5085%, average 73%).
Dysoxic indicators (mainly Bulimina striata) occur
occasionally with low percentages (09%, average
3%). This assemblage occurs predominantly in the
Tortonian portion of Tap 2. The marls are bright
(whitish) and look strongly compacted. Due to the
lack of turbidity flows during the period of deposi-
tion, no bioturbation data are available. Despite of
its high Uvigerina spp. contents, one sample is very
similar to the cluster of the Cibicidoides pachyderma
association, to which it has been allocated (Table 1).
Therefore, the high oxic level is indicated for this
association.
5.4. Globobulimina pyrula association
Dysoxic indicators are the dominant species
(Globobulimina pyrula, Praeglobobulimina affinis,
Chilostomella ovoidea; 4983%, average 68%). Sub-
oxic (1325%, average 20%) and oxic indicators (2
26%, average 12%) are of minor importance. The
assemblages are characterized by very high amounts
of displaced shallow-water foraminifera, indicating
frequent low-density turbidites. Therefore, part of
the oxic indicators may be displaced too. Thus,
this assemblage indicates an oxygen-deficient en-
vironment (dysoxic). Consequently, the associated
turbiditic layers do not show any evidence of bio-
turbation. This association is locally and temporally
very restricted.
6. Water-depth indicator species
As mentioned above, the clustering process did
not distinguished between different water depths.
This is because indicative species for certain bathyal
zones are not frequent enough to influence the
clustering process. This paper follows the water-
depth zonations of Bremer et al. (1980) and Van
Morkhoven et al. (1986): neritic D 0200 m; upper
bathyal D 200600 m; middle bathyal D 6001000
m; lower bathyal D 10002000 m; abyssal D >2000
m. The following species live principally or al-
most exclusively in lower bathyal or abyssal depths:
Uvigerina hispida, 10004000 m, Boersma (1984);
 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
Fontbotia wuellerstorfi, a characteristic species for
the North Atlantic Deep Water (Weston and Murray,
1984) occurs generally >1000 m (several references
in Murray, 1991); Sigmoilopsis celata (Sigmoilop-
sis, 20004000 m, Murray, 1991); Oridorsalis tener
(generally >1000 m, several references in Murray,
1991); and Oridorsalis umbonatus (most frequent
>2000 m, Wright, 1979). Especially U. hispida is
typical for lower bathyal and greater depths. Typical
species of upper bathyal depths (e.g. Planulina arim-
inensis, which has its maximum frequency between
300 and 500 m, Berggren and Haq, 1976) occurs
only sporadically and with very low frequencies.
Real middle bathyal indicators are not mentioned in
the literature and may therefore not exist. A mini-
mum content of 5% of the above-mentioned lower
bathyal species has been regarded as sufficient to
indicate a lower bathyal depth of deposition. The
faunas of all other samples are interpreted as mid-
dle bathyal. Middle to lower bathyal depositional
environments are further supported by planktonic fo-
raminifera contents of 54 to 96% (compare Gibson,
1989). A time correlation of the different forami-
niferal associations with water depth is shown in
Fig. 3.
7. Oxygenation and faunal changes
The most important factor for the foraminiferal
distribution in the investigated area is oxygenation
(possibly in connection with fluctuations of nutri-
ent supply) and therefore it is necessary to briefly
analyse causes for different oxygen levels. Seawater
can dissolve oxygen up to 68.5 ml=l depending
on water temperature, oxygen pressure in the atmo-
sphere and salinity, but this takes place only at or
close to the water surface where photosynthesis and
atmospheric supply maintain a high oxygen level.
Consumption exceeds production below the photic
zone and the oxygen content decreases markedly.
The oxygen level rises again in deep ocean waters
(generally below 1000 m), due to the supply of cold,
relatively dense and oxygen-rich waters by bottom
currents from polar or subpolar regions (e.g. De-
maison and Moore, 1980; Sen Gupta and Machain-
Castillo, 1993). An oxygen minimum zone (OMZ)
149
is developed in the standard ocean water profile
of many modern seas. The oxygen contents of the
OMZ, however, are of wide ranges from <1 to >3
ml=l (Sen Gupta and Machain-Castillo, 1993). An-
other possibility for oxygen depletion is silled basins
with positive water balances, where low salinities
of surface waters causes density stratification with
resultant of stagnant, oxygen-free bottom waters.
Typical examples are the Black Sea (e.g. in Demai-
son and Moore, 1980) or anoxic fjords (Alve, 1990).
A third possibility is coastal upwelling, which may
produce reduced oxygenation by high productivity
and later high oxygen consumption by oxidation
of downsinking organic matter. A zone of reduced
oxygenation (23 ml=l) caused by upwelling is de-
veloped, e.g. off NW Africa between 100 and 700 m
water depth (Lutze and Coulbourn, 1984). However,
upwelling seems to be relatively unlikely at the east-
ern Iberian coast because it normally takes place on
western sides of continents (Demaison and Moore,
1980; Parrish and Curtis, 1982).
If arranged according to their temporal succes-
sion, the investigated sections show a clear tripar-
tition (Fig. 3). The distribution of faunal associa-
tions is subdivided by two marked faunal turnovers.
The first took place during the O. suturalis subzone
(15.215.0 Ma, uppermost Langhian); the sec-
ond during the Gr. siakensis=Gs. obliquus obliquus
subzone (11.711.4 Ma, upper Serravallian). The
intervals before, in between and after the faunal
turnovers were relatively stable periods, at least in
the investigated area. During the Langhian, faunas of
the M. pompilioides=S. reticulata association (dom-
inating) and of the M. pompilioides=C. dutemplei
association (minor) occur in the samples. They rep-
resent predominantly well-oxygenated (high oxic)
environments. The Serravallian is represented by
the almost exclusively occurring M. pompilioides=C.
dutemplei association, indicating reduced oxygena-
tion (low oxic) during this period. Under special
conditions (small sub-basin, section L) oxygen defi-
ciency could develop (G. pyrula association). Well-
oxygenated conditions (high oxic, C. pachyderma
association) were established again in the Tortonian.
The faunal turnovers coincide with other events in
the Mediterranean. The possible reasons which have
caused these changes are discussed below.
 150
H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
Fig. 3. Chronostratigraphic distribution of benthonic foraminiferal associations and inferred depositional depths of the investigated sections and single samples.
 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156 151

8. A palaeoceanographic hypothesis

Gebhardt (1994) proposed that there was an oxy-

gen minimum zone along the eastern Iberian coast
during Langhian and Serravallian times, assuming
relatively stable current patterns for the Langhian to
Serravallian interval as well as for the Tortonian, but
with fundamental differences between them. This
was based on the closer relation of the low oxic M.
pompilioides=C. dutemplei association with the mid-
dle bathyal and of the high oxic M. pompilioides=S.
reticulata association with the lower bathyal (Ta-
ble 1). However, such an interpretation neglects the
results of other studies, in particular those of stable
isotopes, and underemphasizes the faunal turnover at
the LanghianSerravallian boundary (Fig. 3). There-
fore, a re-interpretation appears advisable and a sim-
plified palaeoceanographic model is developed here
(Fig. 4).
The closure of the connection of the Mediter-
ranean to the Indian Ocean during the Early Miocene
(Hsu et al., 1978; Adams et al., 1983; Vergnaud-
Grazzini, 1983) coincides with a worldwide turnover
in the deep-water circulation (Kaiho, 1991). Sluggish
circulation with warm bottom water during the Early
Miocene changed to increased deep-water circula-
tion with cooling of the bottom water in the Middle
Miocene.
From Langhian times onwards, the exclusive con-
nection of the Mediterranean to the world ocean was
to the Atlantic. This remained deep and allowed the
entry of cold water (occurrence of psychrospheric os-
tracods; Benson, 1978) during Langhian time. This is
supported by carbon isotope data (Vergnaud-Grazz-
ini, 1983, 1985) showing the same isotope compo-
sition in Atlantic and western Mediterranean bottom
waters until the end of the Langhian. The entrance
of cold Atlantic water was possible because of the
Fig. 4. Simplified palaeoceanographic model of the western
low position of the Gibraltar sill. Therefore, an
Mediterranean for Langhian, Serravallian and Tortonian intervals.
estuarine circulation can be assumed for Langhian Note that the main triggering factor is the relative movement of
times. the Gibraltar sill. The model is based on interpretations of
At the end of the Langhian (i.e. during O. sutu- Benson (1978, ostracods), Bizon (1985, foraminifera), Gebhardt
ralis subzone, NN5, N9, 15.215.0 Ma) a funda- (1994, benthonic foraminifera), Muller (1985, calcareous nanno-
plankton), Van der Zwaan and Gudjonsson (1986, oxygen and
mental change in the circulation pattern took place,
carbon isotopes), Vergnaud-Grazzini (1983, 1985, carbon iso-
expressed by a faunal turnover among the benthonic topes), Zachariasse and Spaak (1983, planktonic foraminifera),
foraminifera in the investigated area (Fig. 3). This and Weijermars (1991, tectonic events).
was probably caused by a lifting of the Gibraltar
sill as intensive tectonic activity suggests (fold-
152 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
ing, thick turbidite sedimentation in the investi-
gated area; Ott Ott dEstevou et al., 1988; Gebhardt,
1994; widespread discontinuities in the entire Betic
Cordillera; Sanz de Galdeano and Vera, 1992). The
formation of the Arc of Gibraltar (DGibraltar sill)
began at about 15 Ma, together with the uplift of
nappes in the margin of the Alboran Basin (Wei-
jermars, 1991). The restriction of the connection to
the Atlantic may be further supported by sea level
lowering as indicated by Bizon (1985), see also Haq
et al. (1987).
The oxygen isotope signal became distinc-
tively heavier at the LanghianSerravallian bound-
ary (Vergnaud-Grazzini, 1985). This can be inter-
preted as a lower freshwater supply into the Mediter-
ranean as the climate remained tropical (Zachariasse
and Spaak, 1983). One could speculate if chang-
ing configurations of basins and mountain ranges
in the Betic Cordillera (Weijermars, 1991; Sanz
de Galdeano and Vera, 1992) or elsewhere in the
Mediterranean also contributed to this reduced fresh-
water supply or if a more general change towards a
drier climate is the only cause. Less freshwater to
enter the Mediterranean and higher evaporation (sur-
face waters became more saline and therefore denser,
so that they sank to the bottom as it is the case in
the modern Mediterranean) are important factors to
induce a change in the current pattern.
The now starting outflow of more saline Mediter-
ranean water may have contributed to the North
Atlantic Deep Water or its precursors. However, its
amount was unlikely to be sufficiently high to have
been of influence on the formation of the North At-
lantic Deep Water (Vergnaud-Grazzini, 1983, 1985).
The drier climate may be connected with the
worldwide decline in temperature during the Mid-
dle Miocene, generally attributed to the growth of
the Antarctic ice sheet (e.g. Savin, 1977). Antarc-
tic Bottom Water penetrated the deep parts of the
oceans, causing cooling of the bottom waters since
this time.
For the Serravallian in the Mediterranean,
Vergnaud-Grazzini (1983, 1985) interpreted a longer
residence time of the deep water based on carbon
isotopes, causing lower oxygen contents. Also Bi-
zon (1985) reported scarcity or even absence of
benthonic foraminifera faunas in offshore cores of
the western Mediterranean basin, indicating oxygen
deficiency for the Serravallian period. In the investi-
gated area, the high oxic M. pompilioides=S. reticu-
lata association does not appear any more. Instead,
the low oxic M. pompilioides=C. dutemplei associ-
ation strongly dominates; even oxygen deficiency
indicating faunas (G. pyrula association) appear lo-
cally. Thus, a reduced, sluggish circulation caused
lower oxygenation of the bottom waters. Addition-
ally, the interrupted deep-water supply from the At-
lantic resulted in a turn of the circulation towards
an anti-estuarine pattern. Also with the beginning of
the Serravallian, Meulenkamp and Van der Zwaan
(1990) interpreted an increasing nutrient supply for
the deeper (bathyal) waters. An increased stratifica-
tion of the water body has been interpreted by Van
der Zwaan and Gudjonsson (1986), based on carbon
and oxygen isotopes, beginning at about 13.6 Ma.
A distinct cooling started in the late Serravallian
(Zachariasse and Spaak, 1983; based on planktonic
foraminifera), which coincides with a worldwide
drop in temperature.
Renewed tectonic activity all around the Mediter-
ranean (e.g. Bizon, 1985; Chamley et al., 1986, in
Van der Zwaan and Gudjonsson, 1986) affected also
the entrance to the Atlantic (Benson, 1978; Weijer-
mars, 1991; Sanz de Galdeano and Vera, 1992) and
caused a second faunal turnover in the investigated
area (Gebhardt, 1994; this paper) in the late Serraval-
lian. The faunal turnover took place at about 11.7
11.4 Ma (NN7, N14). This coincides with a re-occur-
rence of the oceanic isotope curve in the Mediter-
ranean (Van der Zwaan and Gudjonsson, 1986), in-
dicating an increased exchange with the Atlantic and
a change in the current pattern. This was triggered
most likely by a lowering of the Gibraltar sill,
which caused an intensification of the anti-estuarine
circulation; indicated by higher oxygen content of
the bottom water (this study), similarity of Mediter-
ranean and Atlantic isotope curves (Van der Zwaan
and Gudjonsson, 1986) and stronger inflow of At-
lantic surface water (Muller, 1985). Since that time,
a relatively open circulation similar to todays situ-
ation with a more or less uniform water body (Cita
and Zocchi, 1978) occurred till the final and total
closure of the Mediterranean during the Messinian
by tectonism, on which was probably superimposed
a eustatic lowering of sea level (Van Couvering et
al., 1976).
 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
During the Tortonian, stable environmental con-
ditions with highly oxygenated waters were estab-
lished in the western Mediterranean (Bizon, 1985).
In the investigated area, this is expressed by the only
occurrence of the high oxic C. pachyderma associ-
ation. A stronger inflow of cool surface water from
the Atlantic and absence of temperature fluctuations
is indicated by the nature of calcareous nannofossils
(Muller, 1985) and planktonic foraminifera (Zachari-
asse and Spaak, 1983). Besides the change in the
composition of the foraminiferal associations, the
Tortonian microfaunas show a distinctively lower
diversity (Fisher -indices of 1020 instead of gen-
erally 20 during the Langhian and Serravallian),
lower contents of planktonic foraminifera and a sig-
nificantly increased amount of radiolaria. The higher
amount of radiolaria in the Tortonian samples may
be caused by higher nutrient input from Atlantic
surface waters (see also Brasier, 1995).
Since the Middle Miocene, palaeoenvironmen-
tal differences are indicated between the eastern
and western Mediterranean basins (e.g. Bizon, 1985;
Muller, 1985). The eastern basins experienced less
oxygen supply due to their more restricted configu-
ration (e.g. Cita and Zocchi, 1978; Katz and Thunell,
1984), resulting in less diverse faunas.
Based on recent circulation patterns (Millot,
1987) and the similar arrangement of continental
areas during the Middle Miocene, but much deeper
connections to the ocean than today, an inflow of
deep and cool Atlantic water through the North Betic
Strait, a seaway between the Iberian Meseta in the
north and the rising Betic Cordillera in the south,
and its migration along the northern African margin
is possible for Langhian times. During the Serraval-
lian, middle bathyal intermediate water with reduced
oxygen content may have come from the east and
passed along the southern European coast back into
the Atlantic Ocean, also through the North Betic
Strait. Van der Zwaan and Gudjonsson (1986) inter-
preted an increasingly stratified water body at the
end of the Serravallian in Sicily. Such a stratification
may have developed earlier in the western areas.
They also observed an abrupt end of the stratification
after 10.5 Ma (D10.9 Ma of this paper), which was
only shortly after the benthonic foraminifera faunal
turnover at the Serravallian=Tortonian boundary in
eastern Spain (Gebhardt, 1994; this study).
153
A problem which remains to be solved is the al-
most complete absence of macrofossils in the marl
layers. The only frequent remains of benthonic
macroorganisms are sometimes abundant sponge
spicules. The calcarenitic turbidite layers show bio-
turbation to some extent, but they seem to be
produced exclusively by organisms without hard
parts. According to Demaison and Moore (1980),
the threshold for deposit feeders with hard parts is
about 0.3 ml=l of dissolved oxygen. This is much
below the value which is indicated by the fora-
miniferal faunas. However, more recent reports in-
dicate that severe stress on marine macrobenthos
(hypoxia) may already attain at values of 12 ml=l
in some cases (Boesch and Rabalais, 1991; Tyson
and Pearson, 1991). A further reason may be the
normally low food supply for macroorganisms in
bathyal depths.
9. Conclusions
The following conclusions can be inferred from
the interpretations above.
(1) Oxygenation was the most important factor
for the distribution of benthonic foraminifera in the
Tap Marls.
(2) Two faunal turnovers indicate and coincide
with major changes in the general circulation pat-
terns. The first faunal turnover took place in the
uppermost Langhian (15.215.0 Ma, O. suturalis
subzone, NN5, N9). The second occurred during the
late Serravallian (11.711.4 Ma, Gr. siakensis=Gs.
obliquus obliquus subzone, NN7, N14).
(3) Based on the authors interpretation and ad-
ditional sources, a simplified circulation model for
the western Mediterranean suggests: (a) an estuarine
circulation with relatively high oxygen supply during
the Langhian; (b) a restricted anti-estuarine pattern
with sluggish circulation and resulting reduced oxy-
gen supply until the end of the Serravallian; and (c)
an anti-estuarine circulation similar to todays situa-
tion with high oxygen supply during the Tortonian.
(4) The changes in the circulation pattern were
mainly triggered by tectonic events, in particular
at the connection to the Atlantic Ocean. This was
superimposed by climatic changes and sea level fluc-
tuations.
154 H. Gebhardt / Palaeogeography, Palaeoclimatology, Palaeoecology 145 (1999) 141156
Acknowledgements
I have to express my special gratitude to G.J. Van
der Zwaan, who commented on an earlier version of
this paper as well as to K. Kaiho and J.W. Murray
for their reviews and comments.
Appendix A. List of species mentioned in the text
Following comments from A. Poignant, the names of some
species used by Gebhardt (1994) have been amended. The old
names are given in square brackets.
Ammobaculites humboldti (Reuss) var. latum (Andreae) D Hap-
lophragmium humboldti (Reuss) var. latum Andreae, 1844
Bulimina striata dOrbigny, 1826 [Bulimina costata dOrbigny,
1852]
Chilostomella ovoidea Reuss, 1850
Cibicidoides dutemplei (dOrbigny) D Rotalina dutemplei dOr-
bigny, 1846
Cibicidoides pachyderma (Rzehak) D Truncatulina pachyderma
Rzehak, 1886
Cibicidoides pseudoungerianus (Cushman) D Truncatulina pseu-
doungeriana Cushman, 1922
Cibicidoides subhaidingeri (Parr) D Cibicides subhaidingeri
Parr, 1950
Fontbotia wuellerstorfi (Schwager) D Anomalina wuellerstorfi
Schwager, 1866
Globobulimina pyrula (dOrbigny) D Bulimina pyrula dOrbig-
ny, 1846
Globulina gibba (dOrbigny) D Polymorphina gibba dOrbigny,
1826
Gyroidinoides peramplus (Cushman and Stainforth) D Gyroidina
girardana (Reuss) var. perampla Cushman and Stainforth,
1945
Islandiella laevigata (dOrbigny) D Cassidulina laevigata dOr-
bigny, 1826
Lenticulina stellata (Seguenza) D Robulina stellata Seguenza,
1880
Martinottiella communis (dOrbigny) D Clavulina communis
dOrbigny, 1846
Melonis barleeanus (Williamson) D Nonionina barleeana Wil-
liamson, 1858
Melonis pompilioides (Fichtel and Moll) D Nautilus pompil-
ioides Fichtel and Moll, 1798
Nodosaria longiscata dOrbigny, 1846
Nonion boueanum (dOrbigny) D Nonionina boueana dOrbigny,
1846 [Hanzawaia boueana (dOrbigny)]
Oridorsalis tener (Brady) D Truncatulina tenera Brady, 1884
Oridorsalis umbonatus (Reuss) D Rotalina umbonata Reuss,
1851
Praeglobobulimina affinis (dOrbigny) D Bulimina affinis dOr-
bigny, 1839
Praeglobobulimina pupoides (dOrbigny) D Bulimina pupoides
dOrbigny, 1846
Pullenia bulloides (dOrbigny) D Nonionina bulloides dOrbig-
ny, 1846
Saracenaria italica Defrance, 1824
Sigmoilopsis celata (Costa) D Spiroloculina celata Costa, 1855
Siphonina reticulata (Czjzek) D Rotalina reticulata Czjzek, 1848
Stilostomella consobrina (dOrbigny) var. emaciata (Reuss) D
Dentalina emaciata Reuss, 1851
Stilostomella monilis (Silvestri) var. laevigata (Silvestri) D No-
dosaria monilis Silvestri var. laevigata Silvestri, 1872
Uvigerina barbatula Macfadyen, 1930
Uvigerina hispida Schwager, 1866
Valvulineria complanata (dOrbigny) D Rosalina complanata
dOrbigny, 1846 [Valvulineria bradyana (Fornasini, 1900)]
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