PALAEO-06543; No of Pages 19

Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Benthic foraminifera as palaeoenvironmental indicators during the

last million years in the eastern Mediterranean inner shelf
Simona Avnaim-Katav a,⁎, Ahuva Almogi-Labin b, Amir Sandler b, Dorit Sivan a
a
Department of Maritime Civilizations and the Leon Recanati Institute for Maritime Studies (RIMS), University of Haifa, Mount Carmel, Haifa 31905, Israel
b
Geological Survey of Israel, 30 Malkhe Yisrael, Jerusalem 95501, Israel

a r t i c l e i n f o a b s t r a c t

Article history: Benthic and planktonic foraminifera were studied in inner shelf sedimentary sequences in Haifa Bay, Israel.
Received 23 March 2013 Three boreholes, taken at water depths from 5 to 14 m, spanning about the last 1 Ma, and a set of modern
Received in revised form 11 June 2013 sediment samples used as modern analogue, were utilized for reconstructing palaeoenvironments and
Accepted 12 June 2013
palaeobathymetry. The palaeoenvironmental data were used for assessing tectonic activity of the Carmel
Available online xxxx
Fault, a branch of the Dead Sea transform that bounds the southern side of the bay. Quantitative analyses
Keywords:
showed four biofacies: a fresh to brackish wetland environment, and three progressively deepening marine
Benthic foraminifera biofacies. Relative water depth ranges were estimated using changes in benthic foraminiferal assemblages,
Quaternary presence or absence of Ammonia sp. 1, and the percentage of planktonic foraminifera.
Palaeobathymetry The distribution of the biofacies suggests water depths no deeper than 15 m during most interglacial stages,
Inner shelf water depths to 15–40 m during short transgressive phases of MIS 5.5, 7, 11 and 13, and water depth
Carmel fault reaching 40–80 m in the earliest transgressive phases (MIS 27? and MIS 29?). The environmental conditions
Eastern Mediterranean on both sides of the fault were quite similar along the succession. This suggests that the movements of the
uplifted and down-faulted blocks have more or less coincided since about 1 Ma.
© 2013 Elsevier B.V. All rights reserved.

1. Introduction
Quaternary inner shelf successions located at the sea-land transition
zone would have been frequently affected by glacioeustatic sea-level
fluctuations. Sea-level change is reflected by complexity of the deposi-
tional successions that include hiatuses during lowstands and
palaeoenvironmental changes of the shelf ecosystem during sea-level
highs. Foraminifera have proved to be useful in reconstructing such
palaeoenvironmental oscillations in shallow marine environments, as
the large numbers and diversity enable the group to respond rapidly
to changing ecological parameters (e.g. Mendes et al., 2004; Murray,
2006) and their preservation potential is unparalleled.
The most important controls on benthic foraminiferal distribu-
tion in the Mediterranean Sea and elsewhere are food availability
and dissolved oxygen concentration (e.g. Jorissen et al., 1995; De
Rijk et al., 1999, 2000; Murray, 2001). In shallow water environments
benthic foraminifera assemblages are influenced by additional factors,
including substrate type, temperature and salinity, as for example
recorded in the northern Adriatic Sea, eastern Turkey, the southwestern
Marmara shelf, western Egypt, and off the Israeli coast (Jorissen, 1988;
De Stigter et al., 1998; Basso and Spezzaferri, 2000; Jannink, 2001;
Samir et al., 2003; Hyams-Kaphzan et al., 2008; Phipps et al., 2010).
⁎ Corresponding author. Tel./fax: +972 4 8236819.
E-mail address: simona100@bezeqint.net (S. Avnaim-Katav).
Horton et al. (2007) and Rossi and Horton (2009) showed that the
distribution of living benthic foraminiferal assemblages in shallow
water is highly correlated to water depth. Water depth, in turn, is
connected to substrate type, ratio of siliciclastics to carbonate litholo-
gies (Milker et al., 2009, 2010), oxygen and food availability (Jorissen,
1987; Morigi et al., 2005), or significant changes in salinity, as in del-
taic environments (Mendes et al., 2004).
Quantitative analyses of modern microfauna and sedimentologi-
cal parameters have been used for the palaeobathymetrical and
palaeoenvironmental reconstruction of late Quaternary subsurface
successions in the central Mediterranean (Rossi and Horton, 2009,
and references therein; Carboni et al., 2010, and references therein;
Di Bella and Casieri, 2011).
The abundance, diversity, and composition of benthic foraminiferal
assemblages in the mixed siliciclastic–carbonate ecosystems of the
present southeastern Levantine inner shelf are dependent on substrate
types associated with bathymetry and distance from the Nile cone
(Hyams-Kaphzan et al., 2008). Hyams-Kaphzan et al. (2008) defined
four recent assemblages:
Assemblage A is dominated by the species Ammonia parkinsoniana,
Buccella granulata, Pararotalia calcariformata and some agglutinated
taxa. This assemblage is associated with Nilotic siliciclastic sands,
south of Haifa Bay.
Assemblage B is characterized by the species Ammonia parkinsoniana,
Ammonia tepida, Cribroelphidium poeyanum and Porosononion granosum.

0031-0182/$ – see front matter © 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.palaeo.2013.06.019

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
2 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

This assemblage is typical of Nilotic silty clayey sediments, south of
Haifa Bay.
Assemblage C is characterized by the species Amphistegina lobifera
and Heterostegina depressa and is associated with rocky substrates
in the Levantine carbonate-rich sediments, north of Haifa Bay.
Assemblage D is dominated mainly by Rosalina macropora,
Asterigerinata mamilla, Peneroplis pertusus and Vertebralina striata.
This assemblage is distinctive of carbonate-rich sandy to silty sed-
iments of the northern Levantine shelf.
In this region, foraminiferal assemblage composition has been
influenced by the opening of the Suez Canal in the 19th century, when
Indo-Pacific “Lessepsian” species invaded the eastern Mediterranean
(Por, 1978; Langer and Hottinger, 2000; Hyams et al., 2002). In par-
ticular, Assemblage C is highly dominated by Lessepsian invaders.
Similar assemblages, but lacking the invaders, have been described
in Quaternary coastal sequences along the Israeli shelf (Avital,
2002; Lazar, 2007).
Recent foraminiferal assemblages associated with high and low
energy environments were described by Reinhardt et al. (1994) at

Fig. 1. A. Location of the study area (square) in the eastern Mediterranean, the Dead Sea Transform (DST) and the Carmel–Gilboa Fault (CGF) system following Segev et al.
(2006); B. Morphotectonic map of the Qishon Graben and its surroundings bordered by the Carmel Fault (a part of the CGF system) to the southwest and the Ahihud fault to the
north; C. Bathymetric map (contours at 10 m step down to 110 m depth following Sade et al., 2006) showing location of the studied boreholes (triangles) and the modern analogue
samples in SA and SC transects (circles).

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 3

NW SE A detailed lithology legend

Geomagnetic MSL(m) HB-30

Lithology
Age polarity MIS Age HB-39a
Normal event ka 0 B-3 Sandy silty clay - wetlands

Lithology
Reverse

Lithology
0.3±0.2-
HOLOCENE

1 8.5-8.9 Silty clay - wetlands

10
2 9.9±0.6 -
12.9-13.2
2.2 19.9-20.4
3.1 30.2±2 20 Gravelly sandy clay - palaeosol
5.1 92±9
5.3 88±18 -
5.4 126±9.5 30 Clayey silty sand -
5.5 137±14
139±13
- "Hamra" palaeosol
6.0-6.3 133±14 - 40
6.5-6.6 208±29 Sand
7
7.4 or 216±40- 50
Pringle 8.0 271±48 Sand with aggregates
Falls ? 211±13 - cemented by calcite
B r u n h e s
PLEISTOCENE

227± 8 60
11 397±48 Clayey silty sand
MIS
Excursions?
12 70 event
Excursions? 13 Clayey silty sand with aggregates
13 570±140 80
? cemented by calcite
580±130 21?
?
B/M >780 22? Calcareous sandstone -
M a t u y a m a

25? 90 23? terrestrial (Kurkar)

26? 24? Calcareous sandstone - marine
27? 100
25?
28? Carbonate rocks
110
Jaramillo ~1000 (sandy limestone) - marine
29? H.balthica
26?
<1500-1600
120 Mixed siliciclastic
27? carbonate sediments - marine
Environment
28?
Terrestrial units 130 Sandy silty clay - marine
Marine units 29?
uplifted Carmel down-faulted
Main dating method block block Silty clay - marine
OSL fault
14
C
26 10 Cenomanian dolostone
Al / Be
Bio indicator

Fig. 2. Chronostratigraphy and lithology of the three studied boreholes modified after Avnaim-Katav et al. (2012).

the ancient harbour of Caesarea. Tapiero et al. (2003) showed that qualitative micropalaeontological analyses, were used to reconstruct
diverse foraminiferal assemblages reflect fluctuations in hydrologi- transgression–regression cycles correlated to Pleistocene–Holocene
cal activity attributed to the Nile during the Holocene. fluctuations in global sea levels.
A chronostratigraphically-constrained record of changing envi-
ronmental conditions from the early Pleistocene to the Holocene in
Table 2
the sedimentary sequence of southern Haifa Bay, at the central part The modern analogue samples: sample name, water depth, location, % sand fraction, %
of the Levantine shelf, also showed changes in benthic foraminiferal carbonate content and gear (scuba diving — SD; box corer — BC; R/V Shikmona — S;
assemblages (Avnaim-Katav et al., 2012). Seven marine boreholes Mediterranean Explorer — ME).
were studied, representing parts of the sedimentary record of ap-
Sample Water depth Latitude Longitude Gear % % CaCO3
proximately the last 1 Ma, in which the sedimentary sequence was name (m) (N) (E) N63 μm
found to consist mainly of sand, mixed siliciclastic–carbonate sedi-
SA3 3 32°49.979 35°02.898 SD 98.4 21.4
ments, and clays of marine origin; with sand dunes, palaeosols and SA6 6 32°50.030 35°02.813 SD 97.9 12.0
wetland clays indicating the terrestrial phases. Lithological, sedi- SA10 10 32°50.132 35°02.672 SD 96.6 14.4
mentological, and petrographic changes integrated by a preliminary AS20 20 32°51.501 34°57.874 SD 94.9 20.0
SA30a 30 32°52.210 34°56.580 SD 99.8 87.7
SA40a 40 32°52.447 34°56.100 SD 97.9 91.5
SC3 3 32°53.57 35°04.40 SD 97.5 26.8
SC6 6 32°54.01 35°04.30 SD 93.1 28.0
SC10 10 32°54.02 35°04.18 SD 92.3 39.0
Table 1 SC20a 20 32°54.34 35°02.15 SD 99.0 82.0
The location, water depth and length (m) of the studied boreholes. SC30 31 32°54.42 35°01.16 SD 19.2 19.0
SC40 40 32°55.47 34°56.57 SD 76.0 24.0
Borehole Coordinates Water depth Core length SA50 49 32°53.18 34°55.35 BC, S 74.4 20.1
(m) (m) SA73 73 32°54.73 34°54.30 BC, S 7.0 46.5
Latitude Longitude
SA90 90 32°54.36 34°53.09 BC, S 35.2 8.1
(N) (E)
SA60 60 32°53.533 34°53.689 BC, ME 39.1 29.8
HB-30 32°49.57 34°59.26 5 98.50 SA80 80 32°54.268 34°53.340 BC, ME 2.7 10.9
HB-39a 32°49.59 34°59.49 10.7 120 SA100 103 32°54.409 34°53.183 BC, ME 10.2 8.7
B-3 32°49.15 35°0.48 14.1 121.2 a
Samples excluded from the faunal database.

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
4 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx
The current study mainly focusses on quantitative analyses of the
benthic foraminifera dataset to reconstruct the palaeoenvironments.
A dataset for modern analogues consisting of lithological and
microfaunistic data from the present day Haifa Bay seafloor were
utilized, as a key for interpreting the fossil record. As the study
area was located in a transition zone between the siliciclastic Nilotic
province to the south and the carbonate-rich Levantine province to
the north (e.g., Hyams-Kaphzan et al., 2008), the benthic faunal re-
cords were also used to follow the shifts between these two
sedimentary regimes. The boreholes studied were located on both
sides of the Carmel Fault, a northwest-trending branch of the Dead
Sea Transform fault (Schattner, 2006). The reconstruction of the
palaeobathymetry, palaeoenvironments, and other parameters are
used to evaluate tectonic activity in this area over about the last
1 Ma.
2. Study area
The Mediterranean coast of Israel extends 180 km in a nearly
straight NE–NNE line. The Israeli shelf is 25–30 km wide in the south,
narrowing northward to less than 10 km (Neev, 1965; Almagor and
Hall, 1984). The bulk of the inner shelf sediments are siliciclastics origi-
nating from the Nile River to the south, with a minor contribution from
local sources (e.g. Zviely et al., 2007). The sand is mostly composed of
quartz, with some heavy minerals and small but variable amounts of
biogenic carbonate. Wave-induced longshore currents transport these
sediments predominantly from south to north (Emery and Neev,
1960; Golik, 1993, 1997). Siliciclastics are progressively replaced to-
wards the north by autochthonous carbonate sands in the nearshore
zone (Nir, 1984). Sands predominate in a zone 2.5 to 4 km wide,
extending from the foreshore to ~30 m depth, the approximate position
of the fair-weather wave base. At greater depths, sand is increasingly
replaced by silt and clay (Nir, 1984; Zviely et al., 2007; Almogi-Labin et
al., 2012).
Haifa Bay (Fig. 1) is the most distal locality in which Nilotic mate-
rial can be recognized, and from there to the north, rocky substrates
predominate (Gvirtzman and Buchbinder, 1978; Zviely et al., 2007).
A secondary sediment source, mostly of fines, is the Qishon stream,
which drains into the southern part of Haifa Bay. At present, water
depths in the bay do not exceed 40 m. In the southern part, sand ex-
tends from the shoreline down to a depth of 20 m (Nir, 1980). At
depths greater than 25 m, the sand becomes finer and mixed with
silt and clay (e.g. Zviely et al., 2007). In the central and northern
parts of the bay, Pleistocene calcareous sandstone (kurkar) ridges
occur parallel to the shoreline at 10–25 m depth (Sade et al., 2006),
% >63µm % CaCO3 No. BF/g Species richness diversity index
0 50 100 0 20 40 0 500 1000
0
10
20
Water depth (m)
30
40
50
60
70
80
90
100
110
A B C D
Section SA
Section SC
Fig. 3. Sedimentological and foraminiferal parameters of the modern analogue, Haifa Bay.
Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
forming a hard substrate. Coarse biogenic sediments consisting of
marine carbonate sand and gravel and with a low quartz content
(b10%) fill gullies and channels between the ridges (Almagor et al.,
2000, and references therein).
Haifa Bay was shaped mainly by tectonics. It is the marine exten-
sion of a larger graben structure on land bordered by the normal
Ahihud and Carmel faults to the north and south respectively. The
Carmel Fault, a part of the Carmel–Gilboa Fault system (CGF), is one
of the main branches of the Dead Sea Transform (Schattner, 2006;
Segev et al., 2006). The tectonic activity that caused subsidence of
the Qishon Graben and uplift of Mt. Carmel started in the Miocene,
with an overall vertical displacement estimated at 1500 m. The last
significant uplift of Mt. Carmel of about 300 m occurred during the
early Pliocene (Achmon and Ben-Avraham, 1997; Ben-Gai and
Medvedev, 2005; Zilberman et al., 2010; Gvirtzman et al., 2011).
Other studies have failed to find evidence of activity along this fault
during the last 125,000 years (e.g. Sivan et al., 2001; Medvedev et
al., 2006; Zilberman et al., 2006; Galili et al., 2007). Different
models of the Mediterranean Sea isostatic adjustment concluded
that the induced Holocene isostatic movements have been relatively
small along the coast of Israel (Sivan et al., 2001; Lambeck and
Purcell, 2005).
3. Materials and methods
Continuous cores of the three boreholes HB-30, HB-39a, and B-3,
each of 100–120 m length, were taken in Haifa Bay at water depths
from 5 to 14 m (Figs. 1 and 2, Table 1). The sedimentological character-
istics and the chronological framework of the succession penetrated by
these boreholes were recently described by Avnaim-Katav et al. (2012)
and are briefly summarized in Fig. 2. The chronostratigraphic frame-
work was based on a combination of the dating results (88 dates)
obtained by different methods (optically stimulated luminescence,
14
C, 230Th/234U, cosmogenic isotopes of 26Al/10Be, index fossils and
magnetostratigraphy). The data were correlated to the Waelbroeck et
al. (2002) global sea-level curve and marine isotope stages (MIS). The
sedimentary sequence accumulated over about the last 1 Ma consists
of 21 transgressive–regressive units, representing global sea-level oscil-
lations, with major hiatuses between MIS 5 and MIS 2/3, and between
MIS 11 and MIS 8 (of 80–100 ka and 170 ka duration, respectively). Ter-
restrial beds representing glacial stages include unconsolidated well-
sorted quartz sands alternating with silty clayey sand, representing
palaeosols devoid of fauna, or containing scarce and poorly preserved
faunal remains. The sandy sediments are occasionally indurated into
calcareous sandstones cemented by blocky or sparry mosaic calcite.
Fisher’s-α Evenness
Index Dominance (%) %P
0 20 40 60 80 0 10 20 30 0 1 0 50 100 0 10 20
E F G H
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 5

Some of the terrestrial units include wetland deposits consisting of were used to identify significant groupings, so that the all-retained
organic-rich greyish-brown to black silty clay or sandy silty clay with group had a significant (P b 0.05) internal structure. A ‘similarity per-
rich brackish/fresh-water fauna and plant remains. Conversely, marine centages’ (SIMPER) routine was used in order to identify species that
beds representing interglacial stages consist mainly of loose fine sand contributed most to the similarity within each cluster, as well as to
with relatively well-preserved shallow water faunas. These sands are dissimilarities with other clusters.
occasionally indurated to calcareous sandstone by mainly isopachous The percentage of planktonic foraminifera (%P) within the total
fibrous aragonite cement. Deeper marine units, restricted to the lower foraminiferal assemblage was used as a proxy for palaeobathymetry,
part of the sedimentary sequence, consist of partially cemented mixed following van der Zwaan et al. (1990) and van Hinsbergen et al.
siliciclastic–carbonate sediments, dark silty clays and sandy silty clays. (2005).
All these sediment types are characterized by well-preserved, di-
verse and abundant microfaunas, including benthic and planktonic %P ¼ P=ðP þ B−SÞ  100
foraminifera.
A modern analogue data set, based on 15 recent sediment samples where P = number of planktonic specimens, B = number of benthic
of approximately the top 5 cm, were collected in 2008–2009 from specimens, and S = number of stress-resistant specimens occurring
Haifa Bay along two NW–SE transects (SA and SC), between 3 and in relatively high numbers, including: Bulimina aculeata, Discorbinella
103 m water depth (Fig. 1, Table 2). Three samples of coarse-grain rhodiensis, Fursenkoina acuta, and Nonion fabum (van der Zwaan et al.,
sediments (marked with asterisk in Table 2) were excluded from
the data set, since they had no equivalent in the borehole material.
A. beccarii A. parkinsoniana Ammonia sp. 1 A.tepida
The percentage of N63 μm size fraction was determined by wet siev-
0 5 10 0 40 80 0 15 0 5 10
ing, and CaCO3 content (wt.%) was determined by gasometry.
0
Benthic foraminifera in Recent sediments were stained with Rose
20

Water depth (m)

Bengal (2 g per litre of 90% ethanol), and only the time-averaged
dead assemblage was studied. The use of dead assemblage provides 40
integrated information about the assemblages over a given period of
60
time, whereas the exclusive use of living assemblage represents a mo-
mentary sampling. The occurrence of invading or alien species 80
(Amphistegina lobifera, Hauerina diversa, Heterostegina depressa and 100
Pararotalia calcariformata) introduced to the eastern Mediterranean
120
as part of the Lessepsian migration indicated that the Recent and up-
permost samples in the boreholes were deposited after 1869, when A. mamilla Buccella sp. C. poeyanum D. rhodiensis
the Suez Canal was opened. These species were omitted from the 0 10 20 0 5 10 0 10 20 0 5 10
data set before the statistical analysis. 0
Micropalaeontological analysis was carried out on 158 samples of
20
Water depth (m)

loose and partially indurated sediments from the three boreholes,

sampled every 1 m and at each change in lithology. Partially indurated 40
samples were gently crushed by a press and treated with warm H2O2 to 60
improve disaggregation. Samples of 10–25 g dry weight were washed
80
over a 63 μm sieve, dried, and dry-sieved again. Benthic foraminifera
(N 150 μm) were identified to species level, picked and counted under 100
a binocular microscope, and their state of preservation was evaluated. 120
Some large samples were split into aliquots containing at least 250–
E. crispum P. mediterranensis P. granosum Q. parvula
300 specimens. Samples of terrestrial sediments (e.g. palaeosols and
sand dunes) without microfossils, or with rare and poorly preserved 0 5 10 0 5 0 10 20 0 5 10
0
specimens (with a total abundance b20 individuals per gram dry
sediment (ind./g) were omitted from the quantitative and statistical 20
Water depth (m)

analysis. 40
Taxonomic identification of the benthic foraminifera was based on
Loeblich and Tappan (1987, 1994), Cimerman and Langer (1991), 60
Hottinger et al. (1993), Sgarrella and Moncharmont-Zei (1993), Jones 80
(1994), Reinhardt et al. (1994), Yanko et al. (1994, 1998), Cearreta and
100
Murray (1996), and Reiss et al. (1999). Scanning electron microscope
photographs of key species were taken at the Geological Survey of Israel. 120
Total and relative benthic foraminiferal abundances were deter- R. spinulosa T. bocki T. marioni V. bradyana
mined. Species diversity was expressed as raw diversity (species rich- 0 5 10 0 5 10 0 5 10 0 5 10
ness), Fisher α-index, evenness and dominance (Fisher et al., 1943; 0
Walton, 1964; Buzas and Gibson, 1969; Murray, 1973).
20
Water depth (m)

Q-mode cluster analysis was processed by PRIMER version 6 soft-

ware (Plymouth Routines in Multivariate Ecological Research, UK). 40
The data of the relative abundance of the common benthic species 60
(N3%) and less abundant species which could be grouped into genera,
were double-root transformed in order to down-weight the relative 80
contribution of dominant and abundant species. These transformed 100
abundances were used to build a similarity matrix based on Bray–Curtis
120
similarity, and to reconstruct a dendogram showing hierarchical cluster-
ing (group-average linkage) and a non-metric multi-dimensional scal- Fig. 4. The relative abundance of the common foraminiferal species (N5%) of the mod-
ing (MDS) ordination. ‘Similarity profile’ (SIMPROF) permutation tests ern analogue, Haifa Bay. Full circles are of transect SA and open circles of transect SC.

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
6 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

1990; Edelman-Furstenberg et al., 2001; van Hinsbergen et al., 2005;
Mojtahid et al., 2010).
The genus Ammonia included an unidentified variant with a relatively
large test, with a slightly convex spiral side, whose morphology resembled
the genetic type T13 (Hayward et al., 2004). This variant, referred to
as Ammonia sp. 1, was detected in surface sediments of Haifa Bay at
0–15 m water depth and in surface samples in other localities along the
Israeli coast from the same water depth (Almogi-Labin, personal commu-
nication) and thus was used as a shallow water indicator.
The foraminiferal assemblages (defined by SIMPER) characteriz-
ing the sample clusters (defined by cluster analysis) and their sedi-
mentary characteristics were integrated with their corresponding
palaeoecological parameters (species richness, dominance), %P, and
the occurrence of Ammonia sp. 1 in order to establish biofacies
utilized to interpret the palaeoenvironments and palaeobathymetry.
4. Results
4.1. Modern analogue data set: foraminifera and sedimentology
Fine quartz sands (N90%) occurred between 3 and 30 m water
depth, with the exception of sample SC30 with only 20% sand (Figs. 1
and 3A), in accordance with Hyams-Kaphzan et al. (2008). Sand frac-
tion decreased (~75% sand) at 40–50 m, and at N50 m water depth
the substrate was mainly silty clay (Fig. 3A, Supplementary data No. 1).

Group average
Black lines=significant structure within Transform: Fourth root
(SIMPROF test) Resemblance: S17 Bray Curtis similarity

A
c A. mamilla 11%
b A. parkinsoniana 12% Textularia spp. 7%
A. mamilla 10% P. granosum 7%
Adelosina spp. 6% Adelosina spp. 5% a A. parkinsoniana 57%
Quinqueloculina spp. 5% Triloculina spp. 5% P. granosum 5%
Triloculina spp. 5% D. rhodiensis 5% Quinqueloculina spp. 5%
40 A. beccarii 5% E. crispum 5%
R. spinulosa 5%

60
Similarity

b c a
80

100
SA20

SC30

SC40

SA73

SA80

SA103

SA60

SA50

SA90

SA3

SA10

SC10

SA6

SC3

SC6

Samples

B clusters
a Transform: Fourth root
b Resemblance: S17 Bray Curtis similarity
c
2D Stress: 0.04
SA3

SA60 SA6
SA103 SA20
SC30
SA80 SA50 SC3
SA90 SC6
SA73
SC40
SA10
SC10

Fig. 5. A. Dendrogram of Q-mode cluster analysis of the modern analogue stations, Haifa Bay, using group-average linking of Bray–Curtis similarities calculated on double transform
benthic foraminiferal abundance N3%. Taxa that make significant contributions to the similarity within each cluster (in descending order), based on SIMPER routine, are pointed out
with their raw average abundances. B. Non-metric multi-dimensional scaling (MDS) illustration in two dimensions using the same similarity matrix of the hierarchical clustering
(A). Notice the good separation in the MDS diagram of the three assemblages distinguished in the dendrogram.

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 7

)
(%
x

e
ity -α
de

nc
m

hn es

rs r’s
3
CO

in
s

a
3µ
)

ric eci
es

ve e
y

/g

in
(m

og

Ca

di ish
>6
(m

m
BF

P
Sp
th

ol

Do
F
IS
SL

%
ep

th

.
No
M
M

Li
D

0 50 100 0 50 100 0 1000 0 25 50 0 2 4 6 8 10 0 50 100 0 5

0 5

1
14
10

5.5
20
28
31
30 7
40
44
40 11
49
12
50 55
58

65
13
60
68
71
75 25?
70
81
80 1375
27?

90 97
29?
104
100
A B C D E F G H
Fig. 6. Sedimentological and foraminiferal characteristics of borehole HB-30 (lithological legend in Fig. 2).

The range of CaCO3 concentration was 12–46 wt.% from the shore line to sediment (ind./g) (Fig. 3C). In general, the benthic foraminifera were
73 m and b10 wt.% at greater water depths (Fig. 3B). well-preserved, except at 3 m water depth. Species richness (per sam-
The numerical abundance of the benthic foraminifera varied be- ple) was 28–77, except at 3 m water depth where the low values were
tween less than 100 and up to nearly 1000 specimens per g dry equivalent to Fisher α index value b5 (Fig. 3D–E). High dominance

.
na sp
ia . 1 i um lum
rii on sp sp. hid su um
y a ins a ia a lp u m
pre
s m os ula
log cc ark pid on ell roe isp fab
u
gra
n arv
tho MI
S be p te m cc ib cr de p
Li A. A. A. Am Bu Cr E. E. N. P. Q.
0 5 10 15 0 50 100 0 50 100 0 50 0 10 20 0 10 20 0 10 20 0 10 20 0 10 20 0 5 10 15 0 10 20
0
1
10

20 5.5

30 7
Depth (m)

40 11

50 12

13
60

25?
70

80
27?
90
29?

Fig. 7. Relative abundance (%) of the most common benthic foraminifera species in borehole HB-30 (lithological legend in Fig. 2).

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
8 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

)
(%
x
ity - α

e
de

nc
m

hn es

rs r’s
3
CO

in
s

a
3µ
)

ric eci
es

ve e
y

/g

in
(m

og

Ca

di ish
>6
(m

m
BF

P
Sp
th

ol

Do
F
IS
SL

%
%

%
ep

th

.
No
M
M

Li
D

0 50 100 0 50 100 0 2000 0 40 80 0 2 4 6 8 10 0 50 100 0 5

0 11
1
18
10

31
5.5
20
36
30
7
50
40
54
57 11
50 61

60 12

70 81

88
80 91 13
98
25?
90 101 4132

100 27?

110 121
124
29?
120 131

A B C D E F G H
Fig. 8. Sedimentological and foraminiferal characteristics in borehole HB-39a (lithological legend in Fig. 2).

values of 50–70% that corresponded to a low evenness of 0.1–0.3 oc- between 30 and ~100 m, including Ammonia beccarii, Discorbinella
curred in the sandy sediments at 3–10 m water depth (Fig. 3F–G). The rhodiensis, Elphidium crispum, Reussella spinulosa and Textularia bocki.
dominant species between 3 and 20 m was Ammonia parkinsoniana ac- The value of %P increased with water depth. It was nil at water
companied by Quinqueloculina parvula and Ammonia sp. 1 (Fig. 4). A depths b20 m, whereas between 20 and 40 m it was ~2% (Fig. 3H). At
large number of species reached their maximum relative abundances 50–60 m %P was 3–4% and from 73 to ~100 m it was 10–16% (Fig. 3H).

f.
ian
a
.1 ac na
n p sin nica m ria
gy kin
so
nias ina
ta um yn
e a os
u
nd
a ula rei
be
lo r ida sp Ha germ um n u rv
ho S pa tep mo ca
r
cri fab gra rot pa sc
h
Lit MI A. A. Am C. E. H . N. P. P. Q. T.
0 50 100 0 50 100 0 25 50 0 25 50 0 25 0 25 50 0 5 10 0 5 10 0 5 10 0 5 10 0 25
0
1
10
5.5
20

30 7

40
11
50
Depth (m)

60 12

70
13
80
25?
90

100
27?
110
29?
120

Fig. 9. Relative abundance (%) of the most common benthic foraminifera species in borehole HB-39a (lithological legend in Fig. 2).

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 9

)
(%
x

ce
de
ity α

an
rs s-
in
hn s
s
3
m

CO
)

ric cie
es

ve r’
y

in
(m

og

di she
3µ

F/
(m

P
e
Ca

.B
th

ol

>6

Do
Sp
IS
SL

%
Fi
ep

th

No
%
M

%
M

Li
D

0 50 100 0 50 100 0 2000 0 10 20 30 0 1 2 3 4 0 50 100 0 5

0 14
1
21
10 5.5
28
32
20 7
40
43
30 47

40 11

50 65
69
13
60 76
78
21?
84
70
88
23?
80
8642
98
25?
90 106

100
119
110 125 27?

120 135 29?

A B C D E F G H

Fig. 10. Sedimentological and foraminiferal characteristics in borehole B-3 (lithological legend in Fig. 2).

The Q-mode cluster analysis, based on the 15 recent samples and 29 contributing to each cluster are listed in order of descending average
foraminiferal taxa, distinguished three clusters (Fig. 5A) on a non- relative abundance in Fig. 5A.
metric multi-dimensional scaling (Fig. 5B). The SIMPER routine Cluster a contains samples of sandy substrates in shallow water
highlighted three major sample clusters. The most significant species (b 20 m). The foraminiferal assemblage characterizing this sample cluster

na .
.1 m sp
onia sp s ulu s ula id es um
gy ata ins nia p um res ta res on um os yi
lo fl pa
rk o
cri
s
de
p
ac
u de
p
oe
p b gra
n rad
Litho S
MI A . in A. A mm E. E. F. H. Ne N . fa P. R.
b

0 20 40 0 50 100 0 50 0 5 10 0 5 10 0 5 10 0 5 10 0 5 10 0 20 40 0 10 20 0 10 20
0 1

10 5.5

20 7

30

40 11

50
Depth (m)

60 13
21?
70

80
23?
90 25?

100

110 27?

120 29?

Fig. 11. Relative abundance (%) of the most common benthic foraminifera species in borehole B-3 (lithological legend in Fig. 2).

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
10 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

Group average
A A. parkinsoniana 73% Transform: Fourth root
Aubignyna sp.4% D A. parkinsoniana 56% Resemblance: S17 Bray Curtis similarity
20 A. tepida 14% B. aculeata 3%
E. depressulum 2%
C. auriculus 3% CIII A. parkinsoniana 56%
N. fabum 6% B A. parkinsoniana 85% E. crispum 5%
P. granosum 4% R. macropora 1%
1 CII A. parkinsoniana 37% N. terquemi 1%
E. advenum 2%
Quinqueloculina spp. 12%
40 Adelosina spp. 8%
CI A. parkinsoniana 62%
Triloculina spp. 4%
E. advenum 4%
3 P. granosum 3%
4 R. bradyi 3%
2
3b 3a 3c
Similarity

60

80

100
B3-117.4
B3-51.6
HB39a-49
HB30-47
HB39a-51.6
HB39a-53
HB30-49.4
HB39a-55
HB39a-58
HB30-43.5
HB39a-57.4
HB39a-47.5
HB39a-48.5
HB30-44
HB39a-65
HB39a-65.8
HB39a-22
HB30-93.2
HB30-79.5
HB39a-93
HB39a-95
B3-121.2
HB39a-79.6
HB39a-101.8
HB39a-91.1
HB39a-91.7
B3-63.8
B3-89.6
HB30-56.9
HB39a-73.5
HB39a-70.6
HB39a-77.6
B3-98.5
B3-41.4
B3-54.9
B3-33.6
B3-60.3
HB39a-87
HB39a-11
B3-38.3
B3-74.3
B3-49
HB30-7.9
HB30-4.9
HB39a-0
HB39a-6
HB39a-98.6
HB39a-17
HB39a-102
HB39a-107
HB39a-14
HB39a-69.2
HB39a-90.5
B3-105
B3-66.8
B3-114
B3-84.9
B3-91.1
B3-45.9
HB39a-43
HB30-53.7
HB39a-98
HB39a-110
B3-21.8
HB30-30
HB39a-31
HB39a-32.5
HB39a-34
HB30-22.5
HB30-27
HB39a-37
B3-22.3
HB30-33
HB39a-28
HB39a-35.5
B3-20.3
B3-109.4
HB39a-29.5
HB39a-19
HB39a-20.5
HB30-16
HB30-83.1
HB30-94.8
B3-9
HB39a-118.3
HB39a-96
HB39a-112.9
HB39a-114.6
Samples

Fig. 12. Dendrogram of Q-mode cluster analysis of the 88 samples from Haifa Bay boreholes HB-30, HB-39a and B-3, using group-average linking of Bray–Curtis similarities calcu-
lated on double transform benthic foraminiferal abundance N3%. Taxa that make significant contributions (in descending order) to the similarity within each cluster, based on
SIMPER routine (Supplementary data No. 6), are shown, with their raw average abundances (Supplementary data No. 5).

is dominated by Ammonia parkinsoniana (57%), Porosononion granosum Quinqueloculina spp., and Triloculina spp. Cluster c contains fine
and Quinqueloculina spp. Cluster b consists of samples at water sediment samples from water depths between 40 and 103 m, char-
depths from 20 to 30 m, which are characterized by A. parkinsoniana acterized by A. mamilla (11%), Textularia spp., P. granosum, various
(12%), Asterigerinata mamilla and miliolids, such as Adelosina spp., miliolids, Discorbinella rhodiensis and some other species (Fig. 5A).

4.2. The fossil record

Transform: Fourth root
Resemblance: S17 Bray Curtis similarity
Organic-rich greyish-brown to black silty clay or sandy silty clay
2D Stress: 0.17
with brackish/fresh-water faunas occur in the middle part of the
B3-117.4 sequence. CaCO3 concentrations in these wetland sediments are
4–81%.
The marine sediments consist mainly of yellowish loose fine quartz
sand (N 70%), and occasionally silty clayey sand, sand aggregates or
indurated to calcareous sandstone. These sediments have variable
CaCO3 concentrations, and contain relatively well-preserved faunas
(e.g. Figs. 2 and 6).
clusters
1 4.2.1. Borehole HB-30
2 Borehole HB-30 was drilled at the uplifted side of the Carmel Fault.
3a
3b Its lowermost part was composed of Cenomanian dolostone (Fig. 2).
3c
4
Foraminiferal abundance was variable and relatively low in the
HB39a-22 upper two-thirds of the core, where it never exceeded 350 ind./g. In
Similarity
50 the middle fine-grained unit (of glacial MIS 12), the abundance varied
between 0 and 100 ind./g. (Fig. 6D), whereas the maximum value of
Fig. 13. Non-metric multi-dimensional scaling (MDS) illustration in two dimensions 1375 ind./g occurred in the lower carbonate enriched part, below
computed using the same similarity matrix of the hierarchical clustering (Fig. 12). 79 m (MIS 27?). A total of 126 benthic foraminifera taxa were identi-
The illustration shows relatively good separation of the four assemblages. Two outliers
are indicated by sample name. Partial overlaps between assemblages 3 (lower left) and
fied (Supplementary data No. 2) and species richness (per sample)
2 (centre) and between assemblages 3 and 4 (upper left) are notable, while assem- varied between 1 and 46. High species richness values of 24–46,
blage 1 (upper right) is well separated from the other assemblages. equivalent to maximum Fisher α index values of 3.6–6.7, occurred

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 11

Geomagnetic Borehole HB-30

Age

Lithology
Age polarity MIS ka MSL(m) Borehole HB-39a
Sample clusters
Normal 0 Borehole B-3

Lithology
Reverse 1 2 3 4 Sample clusters

Lithology
0.3±0.2- Sample clusters
HOLOCENE 1 8.5-8.9 1 2 3 4 MIS
10 1 2 3 4
2 9.9±0.6- 1
20 3 2
2.2 19.9-20.4 5.4-5.3
3.1 30.2±2
5.1 92±9 30
5.5
5.3 88±18- 6
5.4 126±9.5
137±14- 7
5.5 40
139±13 8 or 7.4
6.0-6.3 133±14-
B r u n h e s

50
6.5-6.6 208±29
11
PLEISTOCENE

60
7
7.4 or 216±40- 12
Pringle 8.0 271±48 70
Falls 211±13 - 13
227± 8
80 11 ?
11 397±48
13 21?
12 90 22?
15
23?
13 560±140
100 17 24?
B/M ? 580±130 25?
Matuyama

>780
25?
110
26?
27?
26?
28? 120
~1000 27?
Jaramillo 29? H.balthica 28?
<1500-1600 Carmel
fault 29?

Fig. 14. The distribution of the four sample clusters (Fig. 12) against the lithology and chronostratigraphy of the three studied boreholes (lithological and depositional environments
legend in Fig. 2).

in few sandy and carbonate-rich horizons attributed to MIS 13, 7, 5,
and 1 (Fig. 6E–F). Most samples had moderate dominance values of
30–75%, except for several samples of high values, especially in the
middle fine-grained unit (Fig. 6E–G). The most common species
was Ammonia parkinsoniana, occasionally accompanied by Ammo-
nia sp. 1 (Fig. 7). Ammonia tepida, Cribroelphidium sp., Elphidium
crispum and Elphidium depressulum occurred occasionally in high
numbers in the middle fine-grained unit. Values of 3–5 %P were
found at 83 and 85 m (MSL) in carbonate-rich sediments dated to
~ 1 Ma (Fig. 6H).
4.2.2. Borehole HB-39a
Borehole HB-39a was located on the down-faulted side of the
Carmel Fault, 610 m to the east of borehole HB-30 (Fig 1, Table 1). The
foraminiferal abundance largely varied between 0 and b1000 ind./g,
and was even higher in some fine sediment beds in the lower part
(MIS 27?) (Fig. 8D). A total of 190 benthic foraminifera taxa were identi-
fied (Supplementary data No. 3): their species richness values ranged
from 2 to 61, and Fisher α index values were up to ~10 (Fig. 8D–F). In
the fine-grained sediments of the middle part (MIS 12) species richness
was 2 to 23 (Fig. 8A and E). The most common species were Ammonia

Table 3
Palaeoenvironments and estimated palaeobathymetry reconstructions based on biofacies characterization including species richness; dominance; % Ammonia sp. 1 and %P, and the
sedimentary facies.

Biofacies 1 Biofacies 2 Biofacies 3 Biofacies 4

Foraminiferal assemblage Assemblage A Assemblage B Assemblage C Assemblage CIII Assemblage

(CI and CII) D

Species richness Low Low Moderate to high Moderate to high

Dominance Moderate to high High Low to moderate Low to moderate
% Ammonia sp. 1 Common Common Absent Common Absent Absent Absent Absent
P/B ratio 0% 0% 1–3% 0% 1–3% 1–3% 3–6% 3–6%
Sedimentary facies Silty clay rich with Sand to silty-clayey Sand to silty-clayey sand occasionally with sand Silty-clay
brackish ostracods, sand aggregates, or indurated to calcareous sandstone
seeds and plants or partially cemented mixed siliciclastic–carbonate
remains sediments restricted to the base of the succession
Palaeo-environments/Estimated Inland brackish Marine, Marine, Marine, Marine, Marine, Marine, Marine,
Paleo-bathymetry (m) wetlands 0–15 15–40 0–15 15–40 15–40 40–80 40–80

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
12 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

MSL(m) Sample
Age MIS 0 Lithology clusters % CaCO3 % Ammonia sp. 1 %p Palaeobathymetry
5 -40 -80
1 2 3 4 0 50 100 0 25 50 0 5 0-1 15 40
10
HOLOCENE 1
2 20
5.1
5.4-5.3
5.5 30

6
40
7
50
7.4 or
8.0
11 60
PLEISTOCENE

70
12
80

13 90
?
25? 100
26?

27? 110

28? 120
29?

Fig. 15. Palaeobathymetric reconstruction of borehole HB-39a, based on foraminiferal assemblages, species indicative for palaeobathymetry (Table 3) and depositional environments
(lithological and depositional environments legend in Fig. 2).

parkinsoniana and Porosononion granosum (Fig. 9). Cassidulina carinata
and Nonion fabum occurred mainly in the lower part of the sequence,
generally in finer sediments (MIS 27? and MIS 29?). In carbonate-rich
sediments of the lower part the %P was as high as 6% (Fig. 8H).
4.2.3. Borehole B-3
Borehole B-3 was located 2.5 km to the east of borehole HB-30
(Fig. 1, Table 1). The benthic foraminiferal abundance ranged from
80 to 650 ind./g. High values, exceeding 1000 ind./g, occurred in
sands and silty clayey sands (Fig. 10A–D). Clayey sediments at 51
(MIS 12), 117 and 121 m (MIS 29?) depth yielded relatively low fora-
miniferal abundances. Ninety taxa were identified (Supplementary
data No. 4) with a relatively low species richness range of 3 to 28.
The highest species richness value was equivalent to maximum Fisher
α index value of 3.8 (Fig. 10D–F). The most common species was
Ammonia parkinsoniana, followed by Elphidium crispum, E. depressulum,
and Porosononion granosum (Fig. 11). Fursenkoina acuta and Nonion
fabum occurred mainly in the lowermost part of the sequence in finer
sediments (MIS 29?) (Fig. 11). The %P was extremely low, apart from
horizons at 21, 91 and 105 m where the values were 3–4% (Fig. 10H).
4.3. Results of the statistical analyses
The Q-mode cluster analysis, based on 88 samples and 38 forami-
niferal taxa from the three boreholes, illustrated four distinct clusters
and two outliers (samples B3-117.4 and HB39a-22) (Fig. 12, Supple-
mentary data No. 5). The fields of the four main sample clusters on
a multi-dimensional scaling (MDS) diagram are separated. Some
overlaps, however, were observed between clusters 2 and 3, and to
a lesser extent between clusters 3 and 4 (Fig. 13). The two outliers
identified in the dendrogram (Fig. 12) are also shown in Fig. 13.
The distribution of the four sample clusters along the studied
boreholes was similar (Fig. 14). Each sample cluster is characterized
by a rather specific faunal assemblage (referred to as A, B, C, and D),
based on the SIMPER routine (Supplementary data No. 6). The most
significant species in descending order of contribution are listed in
the upper part of Fig. 12 along with their raw average percentage
(Supplementary data Nos. 5 and 6).
Cluster 1 contains mainly samples of borehole HB-39a, from 65.8
to 47.5 m, samples of borehole HB-30, ranging between 49.4 and
43.5 m, and one sample, 51.6 m, of borehole B-3. These samples
occur in the middle part of the sedimentary successions (MIS 12)
and consist of fine-grained sediments. The foraminiferal assemblage
A that characterizes the samples of this cluster is dominated by
Ammonia parkinsoniana (73% of the assemblage on average) accom-
panied by the species Aubignyna sp., Ammonia tepida and some
miliolid species, such as Triloculina spp. Cribroelphidium williamsoni
occurred only in this assemblage and only in borehole HB-30.
Cluster 2 has mostly samples of borehole B-3, from 98.5 to 33.6 m
and one sample, 56.9 m, of borehole HB-30 (all correspond to MIS 11,
13, 21?, 23? and 25?). It has also samples of borehole HB-39a, be-
tween 87 and 73.5 m, that correspond to MIS 12, 13 and 25?, and
also sample 11 m that corresponds to MIS 5.1. The samples of this
cluster consist of loose fine sand and silty clayey sands that are occa-
sionally indurated to calcareous sandstone. This sample cluster is
characterized by a foraminiferal assemblage B dominated by Ammonia
parkinsoniana (85%), Porosononion granosum and Elphidium spp.
Cluster 3 has samples at intervals of borehole HB-39a: 118.3–96 m,
70–69.25 m, 37–28 m, 20.5–14 m, and 6–0 m; of borehole HB-30:

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 13

Foraminiferal assemblage A

1a 1b 2a 2b 3

Foraminiferal assemblage B

4a 4b 5a 5b

5c 5d 5e 6 7

Plate 1. All scale bars are 200 μm.

Foraminiferal assemblage A.
Foraminiferal assemblage B.
(1a) Aubignyna sp., spiral side, (1b) umbilical side (HB-39a, 48. 5–48.55 m). (2a) Ammonia tepida Cushman, spiral side, (2b) umbilical side
(HB-39a, 48.5–48. 55 m). (3) Cribroelphidium williamsoni (Haynes), side view (HB-30, 43.5–43.6 m).
(4a) Ammonia parkinsoniana (d'Orbigny), spiral side, (4b) umbilical side, (HB-30, 1.95–2. 5 m). (5a) Ammonia sp. 1 (Ammonia cf. A. “T13”
Hayward et al., 2004), spiral side, (5b) umbilical side, (5c) lateral view (HB-39a, 0–1.5 m), (5d) spiral side. (5e) umbilical side (SA 6).
(6) Elphidium advenum (Cushman), side view (HB-39a, 17.0–17.45 m). (7) Porosononion granosum (d'Orbigny), side view (HB-39a,
2.5–3.0 m).

94.8 m, 83.1 m, 33–16.5 m, and 7.9–4.9 m; and of borehole B-3: 114–
105 m, 91.1 m, 84.9 m, 66.8 m, 45.95 m, 22.3–20.3 m, and 9 m. These
samples consist of sand to silty clayey sand with aggregates and
partially cemented mixed siliciclastic–carbonate sediments at the
base of the sedimentary successions. Cluster 3 is divided into three
sub-clusters. The foraminiferal assemblage CI that characterizes the
samples of sub-cluster 3a includes Ammonia parkinsoniana (62%), ac-
companied by Elphidium advenum, Porosononion granosum and Rosalina
bradyi. The foraminiferal assemblage CIII that characterizes the samples
of sub-cluster 3c includes A. parkinsoniana (56%), E. crispum, Rosalina
macropora and Neoconorbina terquemi. Sub-cluster 3b is characterized
by A. parkinsoniana (37%), Quinqueloculina spp., Adelosina spp. and
Triloculina spp. The foraminiferal assemblages characterizing the sam-
ples of the three sub-clusters 3a, 3b and 3c have minor additional con-
tribution of diverse species such as Cibicides spp., E. depressulum and
Asterigerinata mamilla (Supplementary data No. 6).
Cluster 4 has mainly samples of borehole HB-39a: 101.8 m, the
interval from 95 to 91.15 m and 79.6 m, samples of borehole HB-30:

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
14 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx

93.2 m and 79.5 m, and sample 121.2 m of borehole B-3. These samples
consist of dark silty clay, or partially cemented mixed siliciclastic–
carbonate sediments restricted to the base of the sedimentary successions
(mainly MIS 27? and MIS 29?). The foraminiferal assemblage D that
characterizes the samples of this cluster has Ammonia parkinsoniana
(56%), Bulimina aculeata, Elphidium depressulum, Nonion fabum, and
Cassidulina carinata. The less abundant species Cancris auriculus and
Ammonia beccarii are part of the foraminiferal assemblage D and CIII
(Supplementary data No. 6). The outlier B3-117.4 has the closest simi-
larity with cluster 4 (Figs. 12 and 13). The foraminiferal assemblage
that characterizes this sample differs from assemblage D by a
high contribution of Ammonia inflata (39%) and N. fabum (36%),
whereas A. parkinsoniana contributes only 11%, and Fursenkoina
acuta, B. aculeata and C. carinata each contributed less than 8%
(Supplementary data No. 5).
5. Discussion
It has been shown that the Recent foraminiferal distribution in the
southeastern Mediterranean shelf (0–40 m) was influenced mainly by
substrate types associated with depth and distance from the Nilotic
source (Hyams-Kaphzan et al., 2008). The complex environmental mo-
saic included siliciclastic and carbonate substrates, as well as coarse to
fine sediments.
The fossil sample clusters of the current study and the foraminiferal
assemblages that characterize them integrated with their correspond-
ing palaeoecological parameters (species richness, dominance), %P,
and the occurrence of Ammonia sp. 1 were used to recognize three ma-
rine and one brackish biofacies. The marine biofacies correspond to the
various sedimentary facies of the cluster samples (Table 3) and are gen-
erally related to depth ranges of the modern analogue. The four
biofacies were used to reconstruct the palaeoenvironments, including
the palaeobathymetry (Table 3, Figs. 14 and 15).
5.1. Palaeoenvironments
5.1.1. Biofacies 1 — inland brackish wetlands
Biofacies 1 corresponds to sample cluster 1, and is characterized
by assemblage A (Plate 1). The samples in this biofacies have low
numerical abundance and species diversity values and moderate to
high dominance. These samples consist of clayey and occasionally
sandy silty clayey sediments, restricted to MIS 12 (Figs. 14, 6 and 8).
The same species of foraminiferal assemblage A are common in
Recent and late Quaternary sediments of brackish marginal Mediter-
ranean environments (e.g. Murray, 2006; Carboni et al., 2010, and
references therein). Moreover, these species were found living in
brackish water bodies (3–5 psu) and in Quaternary brackish sedi-
ments in Israel (Almogi-Labin et al., 1995; Flako-Zaritsky et al.,
2011; Sivan et al., 2011). Ammonia tepida, the dominant species,
is common in organic-rich sediments (e.g. Poag, 1978; Goldstein
and Moodley, 1993). This is in accord with the sample cluster of
this biofacies where flora remains and fresh to brackish ostracods
were encountered, indicating wetland palaeoenvironments. The outlier
clayey sample HB-39a at 22 m depth (Figs. 13 and 14) is dominated by
Haynesina cf. H. germanica (58%), which is also known from brackish,
marginal environments in the Mediterranean (e.g. Murray, 2006;
Carboni et al., 2010, and references therein). In tidal-influenced coasts
this species is characteristic of the upper parts of the tidal flats and
lower salt marsh environments where it is adapted to high seasonal
fluctuations in salinity (Armynot du Chatelet et al., 2009). Nevertheless,
this sample had similar ecological parameter values to this biofacies
(Table 3, Supplementary data No. 3). Accordingly, it may also represent
a brackish water body during the transition from MIS 6 to 5 (termina-
tion II). A certain marine influence, however, is indicated by a low num-
ber of marine species, such as Rosalina macropora, Elphidium spp., and
Quinqueloculina spp. (Supplementary data No. 5 and 6).
5.1.2. Biofacies 2–4 — marine environments
Biofacies 2 corresponds to sample cluster 2, and is characterized by
foraminiferal assemblage B (Plate 1) with low diversity and high
dominance values. The samples consist of loose fine sand and occa-
sionally silty clayey sands. Ammonia parkinsoniana, the dominant spe-
cies, is abundant in the modern analogue of Haifa Bay (Figs. 3 and 4)
and in quartz-rich sediments of less than 30 m water depth along the
Nilotic province (Hyams-Kaphzan et al., 2008, and also Reinhardt et
al., 1994; Yanko et al., 1994; Avital, 2002; Tapiero, 2002). Assemblage
B was comparable to shallow water (b 25 m) assemblages reported
from the southwestern Sea of Marmara that were influenced sig-
nificantly by temperature and salinity (compare also with sub-
cluster C3 in Phipps et al., 2010). Elphidium spp., and especially
Porosononion granosum also appeared in the foraminiferal assem-
blages characterizing clusters 3 and 4. Their presence in sandy to
silty clayey substrates suggested that they were capable of adapting
to variable amounts of food resources (Jorissen, 1988; Jannink,
2001; Avital, 2002; Hyams-Kaphzan et al., 2008; Fig. 4). The highest
abundance and diversity values of these species have been found to
coincide with finer sediments of higher organic matter content. Am-
monia parkinsoniana and A. tepida were dominant during intergla-
cials MIS 5 and 1, and especially in the last 5500 years, reflecting
decreased Nilotic input into the Israeli coast (Avital, 2002; Tapiero,
2002; and compare with Kallel et al., 1997; Calvert and Fontugne,
2001). The foraminiferal composition characterizing the samples of
biofacies 2 is associated with extremely low %P (0–3%), which ac-
cording to the modern analogue indicates b 40 m water depth
(Fig. 3H). In the modern analogue Ammonia sp. 1 is common at
water depth from 0 to 15 m (Fig. 4), and its presence, together
with lack of planktonic foraminifera (Figs. 3H and 4), is used to sub-
divide this range into water depths of 0–15 m and 15–40 m
(Table 3).
Biofacies 3 corresponds to sample cluster 3 and is characterized,
in general, by moderate to high species diversity and low to mod-
erate dominance values. The foraminiferal assemblages (mainly CI
and CIII) characterizing the samples of sub-clusters 3a and 3c (and
to a lesser extent sub-cluster 3b) include epiphytic species as
Rosalina spp (e.g. R. bradyi and R. macropora), Elphidium crispum,
Neoconorbina terquemi and Asterigerinata mamilla (Fig. 12, Plate 2,

Plate 2. All scale bars are 200 μm, except Figs. 1–2, 6b, 8, 11, 14, 500 μm, Fig. 18, 1000 μm.

Foraminiferal assemblage C.
Foraminiferal assemblage D.
(1) Adelosina intricata (Terquem), side view (HB-39a, 6.0–6.45 m). (2) Adelosina laevigata d'Orbigny, side view (HB-39a, 0–1. 5 m).
(3) Quinqueloculina disparilis d'Orbigny, side view (HB-39a, 98.0–98.25 m). (4) Triloculina schreiberiana d'Orbigny, apertural view
(HB-39a, 6.0–6.45 m). (5) Neoconorbina terquemi (Rzehak), spiral side (HB-39a, 96.0–96.4 m). (6a) Rosalina bradyi Cushman, lateral
view (HB-39a, 31.0–31.43 m). (6b) umbilical side (HB-39a, 98.6–99.3 m). (7) Rosalina macropora (Hofker), spiral side (HB-39a, 98.6–
99.3 m). (8) Lobatula lobatula (Walker and Jacob), spiral side (HB-39a, 91.15–91.2 m). (9) Planorbulina mediterranensis d'Orbigny, attached
spiral side (HB-39a, 0–1.5 m). (10) Asterigerinata mamilla (Williamson), spiral side (HB-39a, 6.0–6.45 m). (11) Elphidium crispum (Linnaeus),
side view (HB-39a, 98.0–98.25 m).
(12) Cassidulina carinata Silvestri, umbilical view (HB-39a, 95.0–95.05 m). (13) Bulimina aculeata d'Orbigny, side view (B-3, 117.4 m).
(14) Fursenkoina acuta (d'Orbigny), side view (HB-39a, 114.6–114.7 m). (15) Cancris auriculus (Fichtel and Moll), spiral side (HB-39a,
96.0–96.4 m). (16) Discorbinella rhodiensis (Terquem), spiral side (HB-39a, 91.7–92.0 m). (17) Nonion fabum (Fichtel and Moll), side view
(B-3, 121.2). (18) Ammonia beccarii (Linnaeus) umbilical side (HB-39a, 6.0–6.45 m). (19) Elphidium depressulum Cushman, side view
(HB-39a, 98.6–99.3 m).

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 15

Foraminiferal assemblage C

1 2 3 4

5 6a 6b 7

8 9 10 11

Foraminiferal assemblage D

12 13 14 15

16 17 18 19

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
16 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx
Supplementary data No. 6). These epiphytes are known to be associ-
ated with carbonate-rich sediments (e.g. Murray, 1973; Jorissen,
1987; Kitazato, 1988; Langer, 1993). In the modern analogue they
occurred mainly between 20 and 100 m on sandy and finer substrate
(Fig. 4, Supplementary data No. 1). Foraminiferal assemblages CI and
CIII resemble those recorded in a carbonate unit off Ashqelon that accu-
mulated during the middle Pleistocene (Lazar, 2007). It is also very sim-
ilar to the modern assemblages west of the Nile Delta (Samir et al.,
2003), in Iskenderun Bay, southern Turkey (Basso and Spezzaferri,
2000) and in the northern Adriatic Sea on carbonate-rich sandy plat-
form at water depth from 20 to 100 m (Rossi and Horton, 2009).
Neoconorbina terquemi, Planorbulina mediterranensis, A. mamilla and
R. macropora are common in the carbonate sandy silty sediments
north of Haifa Bay, at water depths of about 30 m (Lazar, 2007;
Hyams-Kaphzan et al., 2008). Foraminiferal assemblage CII, charac-
terizing the samples of sub-cluster 3b, is the only one that contains
abundant miliolids and less frequent epiphytes which characterized
the two other sub-clusters, (Fig. 12, Plate 2, Supplementary data No.
6). The miliolid taxa such as Quinqueloculina species were found to live on
carbonate-rich rocky substrates, whereas Adelosina and Triloculina species
were found mainly on fine-grained sediments (Hyams-Kaphzan et al.,
2008). Most miliolid species (Plate 2) lived in Haifa Bay down to 100 m
on sandy to finer substrate, and showed maximum abundance values at
water depth from 20 to 30 m (Figs. 4 and 5).
In general, the foraminiferal species characterizing the samples of
biofacies 3 are associated with a wide depth range, from the coastline
down to 100 m depth. The combined characteristics of biofacies 3
enabled estimation of the palaeobathymetry (Table 3). The samples
of sub-clusters 3a and 3b consisted of sand to silty clayey sand with
aggregates and of assemblages CI and CII. These assemblages have
extremely low %P alternating from 0% to 1–3%, and presence or
absence of the shallow water species Ammonia sp. 1, respectively.
The reconstructed palaeobathymetry shows two intervals; one ranges
from 0 to 15 m and the other from 15 to 40 m, based on the criteria
that were applied to biofacies 2 (Table 3). The distribution of biofacies
2 and 3 (mainly foraminiferal assemblages CI and CII) indicated depo-
sition in depths from the coastline down to 40 m. The deeper water
depth interval from 15 to 40 m was deposited during interglacial
transgressions MIS 5.5, 7, 11 and 13 (Figs. 14 and 15). Samples of
sub-cluster 3c consisted of partially cemented mixed siliciclastic–
carbonate sediments restricted mainly to the base of the sedimentary
successions and by a foraminiferal assemblage CIII associated with
lack of Ammonia sp. 1, and accompanied by %P alternating between
2–3% and 3–6%. These parameters indicate that during the transgres-
sions of MIS 29? and 27? water depths changed from 15–40 m to
40–80 m, respectively (Fig. 15).
Biofacies 4 corresponds to sample cluster 4, and is characterized by
moderate to high abundance and species richness, low to moderate
dominance values and %P of 3–6% (Table 3). This biofacies is present
exclusively at the base of the sedimentary sequence in MIS 29? and
27? (Fig. 15), and is dominated by dark silty clays, or by partially
cemented mixed siliciclastic–carbonate sediments (Fig. 14). Typical
species in the assemblage (D) characterizing the samples of this
biofacies, apart from Ammonia parkinsoniana (33–83%), were mainly
thin-walled hyaline species, such as Bulimina aculeata, Cancris auriculus,
Cassidulina carinata, Fursenkoina acuta and Nonion fabum (Plate 2).
These species were found deeper than about 70 m in silty clayey sed-
iments in the modern analogue (Supplementary data No. 1). These
shallow to deep infaunal species were associated with fine-grained
sediments where food availability and oxygen concentrations were
found to be the main factors influencing their abundance (e.g.
Jorissen et al., 1992; Goldstein and Corliss, 1994; Altenbach et al.,
1999; De Rijk et al., 1999; Reiss et al., 1999; Mendes et al., 2004).
B. aculeata and N. fabum have been found to be abundant
between 40 and 130 m in the Mediterranean Sea and in the southwestern
Marmara Sea (Mendes et al., 2004; Duchemin et al., 2005; Milker et al.,
Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
2009; Phipps et al., 2010). De Rijk et al. (1999) found B. aculeata
and C. carinata at greater depths, down to 500 m. Ammonia
inflata and the potentially epiphyte Ammonia beccarii (Debenay et al.,
1998) although known to be associated with coarse and sandy substrate
(e.g. Sgarrella and Moncharmont-Zei, 1993) were associated with silty
clayey sediments in Haifa Bay (Figs. 3 and 4). Discorbinella rhodiensis,
found in few samples of cluster 4 and sub-cluster 3c (characterized by
assemblage CIII), is known as a shallow infaunal species that is capable
of adapting to different environmental conditions (e.g. Jorissen, 1988;
Jannink, 2001; Hyams-Kaphzan et al., 2008). This species was found in
early Holocene inner shelf sediments in an assemblage characterized
by high abundance and species diversity values, indicating high produc-
tivity and relative stability of food resources (Tapiero, 2002). D.
rhodiensis lived in Haifa Bay on sandy to finer sediment at depths
from 40 to 100 m (Supplementary data No. 1, Figs. 4 and 5, Plate 2).
Although the species of assemblage 4 have been known from greater
depths, the facts that this assemblage consisted of a relatively low
abundance of Ammonia parkinsoniana, and the %P did not exceed 6%,
constrained the depths to 40–80 m (Fig. 3H).
Biofacies 4 and 3 (assemblage CIII) indicated a transitional sedi-
mentological and faunal regime from the high siliciclastic Nilotic
province to the carbonate-rich Levantine province that prevails
north of the Nile littoral cell.
5.1.3. Biofacies and palaeoenvironment variations with time
The overall distribution of the biofacies along the sequence studied
reveals two major periods (Fig. 16). During MIS 29? and MIS 27? the
reconstructed depths were the deepest in the sequence, and were dom-
inated by biofacies 4 and 3 (assemblages D and CIII, respectively). These
transgressive units consist of unique sedimentary facies of carbonate
rocks and partially indurated mixed siliciclastic–carbonate sediments
interlaced with dark sandy silty clays (Figs. 14 and 16). The overlying
marine units that correspond to MIS 25?, MIS 23? and MIS 21?, pre-
served only in borehole B-3, consist of sands with sand aggregates. In
these units, biofacies 2 and 3 (assemblages CI and CII) were deposited
at depths from 0 to15 m. Since MIS 13, except MIS 12, maximum trans-
gressions are characterized by short deepening intervals to 15–40 m, as
indicated by 1–3% P and the absence of Ammonia sp. 1 (Table 3, Figs. 14
and 16).
The sedimentary sequence (drilled at modern water depths from
5 to 14 m) that corresponds to MIS 12 consists of clayey sediments,
and includes biofacies 1 of wetland setting. It is overlain by three sed-
imentary cycles, which reflect global sea-level oscillations over about
the last 400 ka (Figs. 14 and 16). The sediments of transgressive
phases MIS 11, 7 and 5.5 are sandy. They were deposited in shallow
water with short-term deepening phases (b 40 m), as indicated by
the occurrences of biofacies, 2 and 3 (assemblages CI and CII)
(Figs. 14 and 16). The upper sandy unit of the last transgression,
MIS 1, was deposited at depths from 0 to 15 m, similarly to the
recent water depth as indicated by the occurrences of biofacies 3
(assemblage CII).
Despite the low resolution presented by the studied record we can
still roughly estimate the sea-level at maximum transgression time
during the last 400 ka. This area was tectonically stable during this
period (e.g. Sivan et al., 2001; Medvedev et al., 2006; Zilberman et
al., 2006; Galili et al., 2007; Avnaim-Katav et al., 2012). During MIS
5.5 the sediments accumulated approximately at − 25 m relative to
MSL, whereas the inferred palaeobathymetry ranged between 15
and 40 m (Fig. 16). Using the maximum palaeobathymetry value
the calculated sea-level was higher than the present by few metres,
in accord with previous studies (Lambeck and Chappell, 2001;
Siddall et al., 2003; Lambeck and Purcell, 2005). During MIS 7 and
MIS 11 maximum sea-level was similarly calculated and was found
to be lower than the present sea-level in accordance with previous
studies for MIS 7 (Bard et al, 2002; Antonioli et al., 2004). However,
the estimate for MIS 11 contradicts observations that indicated that
 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx 17

NW SE
Geomagnetic
Borehole HB-30

Lithology
Age polarity MIS MSL(m) Palaeobathymetry Borehole HB-39a
0 0 Borehole B-3
Normal 0 0 0-15 15-4 40-8 Sea Level (m)

Lithology
Palaeobathymetry

Lithology
Reverse Palaeobathymetry
HOLOCENE 1 0 -15 5-40 0-80
10 0 1 4 0 -15 5-40 0-80 -150 -120 -90 -60 -30 0 30
0 1 4
2 0
1
20 3
2
3.1 0.1
5.1 30 5.5
5.3
5.4
40 0.2
5.5 7
B r u n h e s
PLEISTOCENE

50 0.3
6

60
7 0.4 11

Pringle 7.4 or 70
8.0 0.5 13
Falls
80
11 MIS 21?
0.6
12 90 MIS 23?

13 0.7
100
B/M ? MIS 25?
Matuyama

25? 0.8
110
26? 21
27?
28? 0.9 23
120
MIS 27?
29? 25
Jaramillo Carmel MIS 29?
1
27
fault
29

1.1

Fig. 16. Palaeobathymetric reconstruction of the three boreholes (lithological and depositional environments legend in Fig. 2) compared with the global sea-level record of Miller et
al. (2005). Interglacial stages recorded in the studied boreholes were coloured and indicated by odd numbers.

maximum sea-level exceeded the present one (e.g. Olson and Hearty,
2009 and references therein).
5.2. Implication for tectonic activity
The tectonic activity along the Carmel Fault was estimated in a
previous study (Avnaim-Katav et al., 2012) by stratigraphic correla-
tion between boreholes and the relation to their chronostratigraphic
framework. A vertical displacement of 20 m was suggested on the
main fault, and a further downward displacement of 15 m was recog-
nized to the east, suggesting a two-stepped fault in this area. This
total displacement of 35 m occurred in the Early to Middle Pleisto-
cene, from about 990 to 400 ka, but no later distinct tectonic displace-
ment could be observed.
The reconstructed palaeobathymetry indicates that sea-level
alternated between 0–15 m and 15–40 m during large parts of the
sequence, since MIS 25? (~ 950 ka) (Fig. 16). At the base of the
sequence, during MIS 27? and 29?, the depth range was between
40 and 80 m. Water depths and biofacies changes were recorded
equally on both sides of the Carmel Fault all along the sequence,
and could not improve the previous estimation. As the environ-
mental conditions on both sides of the fault were quite similar, it
is suggested that the movements of the uplifted and down-faulted
blocks have more or less coincided since about 1 Ma.
The major tectonic subsidence of the Qishon Graben started in the
late Miocene, with water depths exceeding 200 m (Zilberman et al.,
2010; Gvirtzman et al., 2011). The lower part of the sequence studied,
which accumulated at depths from 40 to 80 m, represents later stages
of the graben infill. These depths did not follow the global sea-level
curve (e.g. Miller et al., 2005) but were rather a result of keeping up
the accommodation space.
Maximum depth was shallower and quite stable through time,
since MIS 25?, indicating that subsidence rates were balanced by sed-
iment accumulation rates of the Qishon Graben.
6. Conclusions
The fossil benthic foraminiferal record of Haifa Bay supports pre-
vious observations that benthic foraminifera are good indicators for
reconstructing past environments, since they are sensitive to changes in
the environmental conditions, particularly at the sea-land transition zone.
Four distinct sample clusters were defined by cluster analysis in
the subsurface of southern Haifa Bay. Each cluster is characterized by
sedimentary facies and foraminiferal assemblages. These assemblages,
together with their corresponding ecological parameters, %P, occur-
rence of Ammonia sp. 1, and the characteristics of the modern ana-
logue, were used in reconstructing the four biofacies that represent
environmental changes over about the last 1 Ma:
Biofacies 1: dominated by Ammonia parkinsoniana, Aubignyna sp.,
Ammonia tepida, Cribroelphidium williamsoni, Cribroelphidium sp., some
miliolids, fresh to brackish ostracods, and some flora remains. This
biofacies consists of clayey to sandy silty clayey sediments, and represents
mainly fresh to brackish wetland environments during glacial MIS 12.
Biofacies 2 and 3 (mainly foraminiferal assemblages CI and CII)
represent marine environments with depths between 0 and 15 m and
between 15 and 40 m, based on the presence or absence of Ammonia
sp. 1 and %P of 0% or 1–3%, respectively. Ammonia parkinsoniana,
Porosononion granosum and Elphidium species occur in Biofacies 2,

Please cite this article as: Avnaim-Katav, S., et al., Benthic foraminifera as palaeoenvironmental indicators during the last million years in the
eastern Mediterranean inner shelf, Palaeogeography, Palaeoclimatology, Palaeoecology (2013), http://dx.doi.org/10.1016/j.palaeo.2013.06.019
18 S. Avnaim-Katav et al. / Palaeogeography, Palaeoclimatology, Palaeoecology xxx (2013) xxx–xxx
which was typical of quartz-rich sands. In Biofacies 3 typical sediments
were sand to silty clayey sand with aggregates characterized by epi-
phytes species, such as Rosalina spp., Neoconorbina terquemi, Cibicides
spp., Asterigerinata mamilla, Planorbulina mediterranensis, Lobatula
lobatula, and miliolid taxa. Biofacies 2 and 3 are characterized mainly
by assemblages CI and CII that represent marine environments with
shallow depths of 0 to 15 m. They prevailed during interglacial stages
since MIS 25 with short deepening intervals of depths between 15
and 40 m during interglacials MIS 13, 11, 7 and 5.
Biofacies 3 is characterized also by assemblage CIII and consisted of
epiphytes (as in foraminiferal assemblage CI) in mixed siliciclastic–
carbonate-rich sediments. It represents depths between 15 and 40 m
with %P of 2–3%, or between 40 and 80 m with %P of 3–6%.
Biofacies 4 is characterized by assemblage D with deeper water in-
faunal species, such as Bulimina aculeata, Cancris auriculus, Cassidulina
carinata, Fursenkoina acuta, and Nonion fabum. The sedimentary fa-
cies consist of fine-grained sediments, or partially cemented mixed
siliciclastic–carbonate sediments and %P of 3–6%, indicating depths
from 40 to 80 m. Biofacies, 4 and 3 (assemblage CIII) represent
marine environments with alternating depths from 15 to 40 m, and
from 40 to 80 m during MIS 29? and MIS 27?
The environmental conditions on both sides of the Carmel Fault
were quite similar during the last 1 Ma suggesting that the move-
ments of the uplifted and down-faulted blocks have more or less
coincided during this period.
Supplementary data to this article can be found online at http://
dx.doi.org/10.1016/j.palaeo.2013.06.019.
Acknowledgements
This study was carried out with the support of an Earth Sciences
Administration Grant of the Ministry of National Infrastructures, a
Sir Maurice Hatter Research Grant of the Leon Recanati Institute for
Maritime Studies (RIMS), and the University of Haifa Research
Funds. We gratefully acknowledge the technical assistance received
from M. Kitin and M. Dvorachek, Geological Survey of Israel. Thanks
are due to Y. Melamed of the Bar-Ilan University of Ramat-Gan for
identifying the flora remains. We deeply thank G. Schmiedl and the
other anonymous reviewer for their constructive comments and sug-
gestions resulting in a significantly improved manuscript.
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