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Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44

Contents lists available at ScienceDirect

Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o

Climate variability in the Upper Jordan Valley around 0.78 Ma, inferences from
time-series stable isotopes of Viviparidae, supported by mollusc and
plant palaeoecology
B. Spiro a,b,⁎, S. Ashkenazi c, H.K. Mienis c, Y. Melamed d, C. Feibel e, A. Delgado f, A. Starinsky g
a
Institute of Earth Sciences, The Hebrew University, Jerusalem 91904, Israel
b
Department of Mineralogy, Natural History Museum, London SW7 5BD, United Kingdom
c
National Natural History Collections, Berman Bldg., The Hebrew University, Jerusalem, 91904, Israel
d
The Martin (Szusz) Department of Land of Israel Studies and Archaeology, Bar-Ilan University, Ramat Gan 52900, Israel
e
Department of Geological Sciences, Rutgers University, Busch Campus, 610 Taylor Road, Piscataway, NJ 08854-8066, United States
f
Estacion Experimental del Zaidin (CSIC), Granada 18008, Spain
g
Institute of Earth Sciences, The Hebrew University, Jerusalem 91904, Israel

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

Article history: The Early–Middle Pleistocene lacustrine sediments at Gesher Benot Ya'aqov (GBY) in the Upper Jordan Valley
Received 14 August 2007 span ca. 100 Ka across the Matuyama–Brunhes Boundary (MBB) at 780 Ka and oxygen isotope stages (OISs)
Received in revised form 27 July 2009 20 to 18, provide a continuous record of their sedimentary history and prevailing climate. The lacustrine
Accepted 6 August 2009
environment is investigated using ecological and stable isotope characteristics of gastropods belonging to the
Available online 18 August 2009
family Viviparidae supplemented by other molluscan and floral indicators. The sedimentary framework of
the 34 m thick sequence is bounded by channel conglomerates at its base and top, and consists of six
Keywords:
Molluscs
sedimentary cycles. Each one starts with shore sediments (coquina and sand) and ends with deeper water
Jordan Valley sediments (black mud or gray mud).
Early–Middle Pleistocene Time series of δ18O and δ13C values were determined in individual shells belonging to two sympatric genera of
Stable isotopes viviparid gastropods: Viviparus apameae galileae and Bellamya sp. found in the sedimentary cycles 2 through 5.
Palaeoenvironment The total overall range of δ18O in the time series of all the shells is from −10.5‰ to −5.2‰ (VPDB), and the δ13C
Lacustrine sediments from −9.2‰ to −4.8‰ (VPDB). The δ13C values of shells recovered from black mud are lower than those from the
other sediment types suggesting a higher proportion of carbon of organic origin. The δ18O values show only a
weak relation with sediment type. The time series of the individual shells indicate a relative constancy in the
water δ18O and temperature whilst the variation in δ13C could be related to the annual productivity cycle (growth
and decay of aquatic plants) of the lake. The δ13C patterns in the time series and winter rings indicate that the rate
of growth of the individual snails was ca. 15–30 mm/year. The isotopic characteristics of the Viviparid shells from
these beds indicate diverse sedimentary environments and suggest that above the MBB (cycles 2–5) the GBY lake
received constant and isotopically consistent water inflow.
Our integrated data, isotopic and ecological of molluscs, and ecological of plants, indicates that the geophysical-
magnetic phenomenon of MBB coincides with an environmental change from a relatively cold dry climate
towards a warmer temperate subtropical climate in this region. This change coincides with the occurrence of
hominin artifacts, which were found mainly in layers of the sedimentary cycles 3–5.
© 2009 Elsevier B.V. All rights reserved.

1. Introduction carbon and oxygen in biogenic carbonates. Their interpretation is greatly

1.1. The Gesher Benot Ya'aqov site
The reconstruction of the palaeohabitat and palaeoenvironment is a
multidisciplinary endeavour that requires biological and ecological
information on species that have restricted environmental requirements,
and is facilitated by geochemical indicators such as stable isotopes of
⁎ Corresponding author. Institute of Earth Sciences, The Hebrew University,
Jerusalem 91904, Israel.
E-mail address: b.spiro@nhm.ac.uk (B. Spiro).
enhanced by comparison with recent habitats for which the environ-
mental parameters can be directly related to their living flora and fauna
with their particular ecological conditions and the respective geochem-
ical indicators. This study uses this approach to interpret the palaeocli-
mate of the Early–Middle Pleistocene (780 Ka) Acheulian, lakeside site of
Gesher Benot Ya'aqov (GBY) located in the Northern Jordan Valley, Israel,
part of the Syrian–African Rift (Fig. 1). This site is of major importance for
the elucidation of Early–Middle Pleistocene climate in the Middle East
and the hominin “out of Africa” migration. The site contains a well
documented, 34 m thick sedimentary sequence, which crosses the
globally identifiable Matuyama–Brunhes Boundary (MBB) at 780 Ka and

0031-0182/$ – see front matter © 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2009.08.005
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B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44
Fig. 1. Location map of the Gesher Benot Ya'aqov (GBY) site in the Upper Jordan Valley.
has a long time span of ca 100 Ka (Verosub et al., 1998; Goren-Inbar
et al., 2000; Belitzky, 2002; Feibel, 2004). This time span includes the
well-defined climate variations recorded in coeval deep sea
sediments of OIS 18–20 (Bassinot et al., 1994; Feibel, 2004) and the
Antarctic EPICA Dome C ice core record, which as of 2004 provided D/
H records up to 740 Ka (EPICA, 2004). The GBY sequence includes at
least ca 45 discrete waterlogged sedimentary beds containing very
rich and diverse floral and faunal remains (Melamed, 1997; Goren-
Inbar et al., 2002a,b; Melamed, 2003; Feibel, 2004; Rabinovich et al.,
2008). At least 13 of these sedimentary beds are archaeological
horizons with evidence of human habitation and activity with
assemblages of lithic artefacts (Goren-Inbar et al., 2000; Feibel, 2004;
Goren-Inbar et al., 2004) (Fig. 2).
1.2. Fossil Viviparidae in the Upper Jordan Valley
The present study investigates fossil species of the gastropod
family Viviparidae from the diverse mollusc assemblages of GBY. So
far 71 species belonging to 19 families have been identified in the
section (Ashkenazi and Mienis, 2005). In general the prosobranch
33
species of the Viviparidae are relatively large (up to 50–60 mm high)
freshwater snails. The genus Bellamya Jousseaume, 1886, from this
family is known from southern Asia and Africa (Prashad, 1928; Brown,
1980; Sivan et al., 2006). Extant species inhabit rivers, lakes, swamps
and ponds, ranging in water depth from 0.3 to 3 m. Viviparidae were
not reported from marine or brackish water indicating that salinity is
the main factor that limits their occurrence (Prashad, 1928). The
salinity tolerance of recent Viviparidae populations is generally lower
than 0.3–0.7 ppt (Glöer and Meier-Brook, 1998). They prefer a soft
substrate and muddy habitats with aquatic macrophytes and algal
mats (Jokinen et al., 1982). In cold climates they burrow in the
sediment for hibernation (Ribi et al., 1986). They are filter feeders on
diatoms, algae and detritus, but occasionally become carnivores
(Cook, 1949; Browne, 1978). Female Viviparids, carry developing
embryos with shells and give birth to fully developed young. Certain
extant species in North America grow continuously throughout the
year and throughout the life cycle, although the growth rate decreases
during the winter (Jokinen et al., 1982). The life spans of two recent
European Viviparus species V. viviparus (Linnaeus 1758) and
V. contectus (Millet 1813) were found to be 6–7 years (van der
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34 B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44

Fig. 2. Stratigraphy of the individual archaeological trenches at Gesher Benot Ya'aqov (based on Feibel, 2001). Abbreviations for lithology: c — clay, z — silt, s — sand, g/q — gravel or
coquina.

Spoel, 1958). Extinct species of the Viviparidae are found in the
Pleistocene fossil coquina beds along the Jordan Valley and the
Orontes River region (Kinzelbach, 1980; Schütt and Ortal, 1993). The
coquina beds in the Upper Jordan Valley were defined by Picard
(1963) as Viviparus beds of the Benot Ya'akov Formation with the
index fossil Viviparus apameae. The nominal subspecies Viviparus
apameae apameae (Blanckenhorn, 1897) is thought to have originated
in the Orontes River system. During the Miocene it probably reached
the middle sector of the Jordan Valley (Tchernov, 1975), where it
developed into a local subspecies Viviparus apameae galileae Schütt
and Ortal (1993). The study of a continuous core from the Hula Valley,
representing a sequence from the present to 700 Ka, indicates that this
Viviparus disappeared suddenly below the Main Peat (Moshkovitz and
Magaritz, 1987). Peat samples, just above the last Viviparus specimens
in the core holes L1 2 and L2 1 have been dated by the U/Th method to
222 Ka (Kafri et al., 1983). Based on this dating, the extinction of
Viviparus apameae galileae occurred in the Hula Basin at ca 240 Ka
(Moshkovitz and Magaritz, 1987). There is no direct explanation for
the disappearance of Viviparus, it is however accompanied by a
gradual decrease in δ18O values towards the bottom of the core-hole
indicating an (upward) shallow phase of the lake with increased
evaporation (Moshkovitz and Magaritz, 1987).
1.3. Stable isotopes in gastropods and their
palaeoenvironmental interpretation
The δ18O and δ13C values of freshwater molluscan shells have been
used as high-resolution records of environmental conditions and their
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B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44
changes in the aquatic habitat during the period of their growth, and
thus provide a powerful tool in palaeoclimate research. These δ18O
records mainly reflect the δ18O of the water and its temperature. The
temperature dependent fractionation for aragonite precipitation is ca
0.24‰ per degree (Kim and O'Neil, 1987). However 1 °C decrease in
temperature lowers the δ18O of the rain water by 0.69‰ (Dansgaard,
1964) and hence the effect of change in rain water composition is
more important than the temperature of mineral precipitation.
Furthermore for the formation of shells this fractionation may be
affected by kinetic isotope effects due to biological controls on shell
deposition. Isotope profiles were constructed in this study in a
manner similar to those of Abell (1985) and Leng et al. (1999) who
studied African freshwater shells. Abell et al. (1995) who performed
time-series analyses to study seasonality along spirals of recent
freshwater shells in Africa concluded that the relation between the
oxygen isotope composition and environmental parameters was
complex. Various studies highlighted different factors affecting this
complexity. While Leng et al. (1999) reported that the δ18O values in
shells of modern Melanoides tuberculata (Müller 1774) from Lake
Awassa were in equilibrium with the lake water, Shanahan et al.
(2005) observed that gastropods living in nearly constant conditions
showed variations larger than predicted even for Melanoides
tuberculata. Schmitz and Andreasson (2001) showed that the
calculated values of δ18O of the latest increments in gastropod shells
determined according to Grossman and Ku (1986) on the basis of the
isotope composition and temperature of the ambient waters, were
systematically higher than those measured. The difference varied
between 0.3, 0.5 and 1.09‰ for the species Stagnicola palustris Müller
1774 (as Lymnea palustris), and 1.4‰ for Haitia acuta (Dransparnaud
1805) (as Physa acuta). Thus they could calibrate the offset for the
respective species. This comparison implies that there is a consider-
able uncertainty associated with the interpretation of δ18O values of
gastropods with respect to ambient environmental conditions. The
respective differences in temperature could correspond to ca 3 °C (see
also Andreasson and Schmitz, 1996, 1997).
The δ13C values of gastropod shells were found to relate to that of the
Dissolved Inorganic Carbon (DIC) reservoir (Letizia et al., 1997). The
effect of respired (metabolic) carbon was found by Shanahan et al.
(2005) to be small in comparison with the contribution from DIC (2% for
Melanoides). Whilst Geist et al. (2005) suggest that in the freshwater
pearl mussel Margaritifera margaritifera, Linnaeus 1758, this contribu-
tion may be larger. Shanahan et al. (2005) suggest that the interpre-
tation of the stable isotope results requires taking into account both
behavioural and physiological and probably also kinetic factors.
Schmitz and Andreasson (2001) followed a comparative approach
to palaeoenvironmental interpretation of stable isotopes in fossil
gastropods based on high-resolution isotope analysis of time series in
samples of extant related species from similar habitats. This led them
to the identification of the climatic zone on the basis of similarity
between the isotopic characteristics of extant specimens of specific
climatic zones with those of fossil specimens. The time series of extant
species they report include three species of viviparids from the
Mediterranean (V. hellenica (Clessin, 1879), V. fasciatus Müller 1774
and Bellamya polita). The common features are a large change in δ18O
towards the aperture and great variability within the δ13C profiles.
The present study examines two extinct sympatric species of the
Viviparidae family having the same ecology and habitat that were
collected from a single site and represent a time span of ca. 60 Ka.
Therefore the effects of geographic factors (such as latitude, altitude
and the characteristics of the catchment), taxonomy, physiology,
behaviour, ecology and general habitat can be regarded as constant for
these two species over this period.
In view of the various factors affecting the interpretation of the
stable isotope records of gastropods discussed above, the present
study concentrates on the isotopic composition (δ18O and δ13C) and
mainly its variations as indicator of environmental change.
35
2. Aims
The present study aims to interpret the long-term environmental
record reflected in δ18O and δ13C values of the aragonitic Viviparidae
in the Lower–Middle Pleistocene (780 Ka) freshwater lakeside
sediments of Gesher Benot Ya'aqov (GBY). This will be achieved by
time-series sampling and analysis of viviparid shells to assess seasonal
and inter-annual amplitudes of δ18O and δ13C values in order to
understand the yearly and inter-annual changes of ambient water and
productivity. The interpretation based on the isotope analysis will be
compared with available information on the sediment, flora and fauna
of the respective beds for palaeoenvironmental reconstruction of the
lakeside site and the surrounding area. This will contribute to the
analysis of climate and climate change in a pivotal area on the “out of
Africa” migration route with implications on more recent environ-
mental changes in this area.
3. Material and methods
3.1. The stratigraphy at the GBY site
The GBY archaeological site (33°00′30″N, 35°37′30″E) is located
within the GBY embayment, which was formed by the uplift of the
Korazim Saddle in relation to the pull-apart graben of the Hula to the
north (Belitzky, 2002) (Fig. 1). The sedimentary setting is described by
Goren-Inbar et al. (2000) and Feibel (2004).
The series of samples investigated in the present study were taken
from individual well-identified discrete sedimentary beds described by
Feibel (2004) and given in Fig. 2. The section below the MBB was
sampled only from trenches, whilst the overlying sediments were
sampled both from trenches and from larger archaeological excavations.
The whole stratigraphic sequence is bounded by channel fluvial
conglomerates at bottom and top, and was interpreted as constituting
a single first-order Milankovich cycle. This first-order cycle consists of
six second-order cycles reflecting changes within the lacustrine
environment (Feibel, 2004). These six second-order cycles are referred
to here as cycles 1–6. Each one of these cycles (except the top bounding
conglomerate) consists of beach sediments (coarser-grained) overlain
by lake deposits (fine-grained). Two types of off-shore muds occur; gray
mud (oxic) in cycles 1 and 2 and black organic rich mud (anoxic) in
cycles 3–5 (Fig. 2) (Feibel, 2004). The sedimentary features in each cycle
represent shore and deeper water and hence a decrease and increase in
water depth of the lake respectively. Most of the layers are predomi-
nantly carbonate (up to 90%), including both fine-grained micrite and
calcareous shells (Feibel, 2001) indicating that the palaeolake was a
marl lake. The stratigraphic position and sedimentological definition of
the individual samples were given by Feibel (2004). This record of a
regular cycle of water level in the GBY palaeolake suggests that no major
tectonic events disturbed the sedimentary regime during this period
estimated at 100 Ka and therefore the cyclic changes in the
sedimentological characteristics should be ascribed to environmental
changes possibly driven by a change in climate. On the basis of the MBB
these cycles were correlated with the synthetic oxygen isotope
stratigraphy (OIS) of Bassinot et al. (1994) ranging between OIS 20
and 18 (Feibel, 2001, 2004).
4. Mollusc samples
The taxonomy and nomenclature of the fossil Viviparidae
discovered so far in the Levant are debated. Two species were
reported from Israel (Picard, 1965; Tchernov, 1973, 1975; Kafri et al.,
1981; Steinitz and Mor, 1982). Recently, Sivan et al. (2006)
recognized four different species, of which one remains undescribed.
Most recent studies mentioning Viviparids from GBY in detail
(Tchernov, 1973; Schütt and Ortal, 1993, and in a lesser degree Sivan
et al., 2006) were based on non-stratigraphic collected material. In
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36 B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44

concentrate mainly on the Viviparidae and refer to other species

only for discussion. The largest and least damaged Viviparus or Bel-
lamya specimen from each bed was selected for the time-series stable
isotope analysis. The specimen's height and when possible width,
were measured and recorded (in mm) and the number of whorls was
recorded from whole shells (Table 1). In two of the samples the only
representative specimens of Viviparidae were broken parts of the
shell. In the present study we assume that the two species occupied
the same habitat and had the same ecological requirements probably
similar to those of extant representatives of this family.

4.1. Stable isotope analysis

Each specimen was cleaned in an ultrasonic bath. Aliquots for δ18O

Fig. 3. Viviparidae from GBY, a) Viviparus apameae galileae and b) Bellamya sp. scale bar
2 cm.
and δ13C analysis were obtained by drilling with a 2 mm diamond
coated drill. These were taken from most specimens at 2.5–3 mm
intervals starting with the latest growth increment just behind the lip
of the aperture. The first specimen investigated (Bellamya from bed V-
6) was sampled at ca. 1 mm interval. X-ray diffraction analysis (Phillips
1830 Cu Ka radiation) of drilled aliquots showed the indicative trace of
aragonite with no indication for the presence of calcite.
Stable isotope analysis of the drilled aliquots was carried out on a
Finnigan MAT 253 connected to a ‘Kiel device’ (Thermo electron
corporation) at the Stable isotope laboratory UCL London UK. Aliquots of
the NBS 19 standard run along with the samples gave −2.23 ± 0.06‰
(1cr) for δ18O (n = 9) and 1.95 ± 0.02‰ (1cr) for δ13C (n = 9). Results
are reported relative to VPDB with an overall analytical reproducibility
for sample aliquots of less than 0.10‰ and 0.05‰ for δ18O and δ13C
respectively.
5. Results

this way Tchernov (1973: plt. 6, Figs. 1–4) and Schütt and Ortal (1993:
plt. 1, Figs. 1–13) created a composite species, Viviparus apamea
galileae. The material used in the present study belongs to two
morphologically distinguishable species (Fig. 3), which occur within
the GBY sequence either in different layers or sympatric in the same
layer (Ashkenazi and Mienis, 2005). Here we refer to the distinct
forms of the Viviparidae from GBY as Viviparus apameae galileae and
Bellamya species.
Specimens of Viviparus apameae galileae and Bellamya sp. were
sorted from 20 consecutive bulk sediment samples (350–500 g each)
from the well-identified and discrete stratigraphic levels, discerned
down to 10 cm thickness along the excavated sequence of GBY, both
above and below the MBB (Fig. 2). The samples were selected to
represent all sediment types with minimum gaps between two
successive samples. All mollusc specimens in the samples were
identified to the lower possible taxonomic level, and then counted
and recorded (Ashkenazi and Mienis, 2005). In this study we
5.1. The distribution of Viviparid shells in the stratigraphic levels of GBY
A total of 45 distinct beds were identified and sampled in the
sedimentary sequence of GBY shown in Fig. 2. Out of these 20 were
chosen for detailed examination in this Study; 10 were from the
sequence below the MBB (cycle 1) and 10 were from the sequence
overlying it (cycles 2–5). Specimens of Viviparidae were found only in
10 layers overlying the MBB (9 layers and one sample that represents
the contact between two layers) in the interval 12.2 and 26. 6 m above
the base of the section. No Viviparid shells were discovered in the
sediments of cycle 1 and only fragments in cycle 2 (bed II-14) the first
bed above the MBB.
Ten Viviparidae specimens of the genera Viviparus and Bellamya
from these beds were analyzed for δ18O and δ13C (Table 1, Fig. 2).
Eight specimens provided time series of between 7 and 43 serial
aliquots, two samples consisting of single fragments provided only 3
aliquots each. The specimens analyzed were from different sediment

Table 1
Samples of specimens of Viviparus apameae galileae and Bellamya species from GBY sequence examined for time series of δ18O and δ13C isotope data. Stratigraphic levels, cycles and
sediment type are based on Feibel (2004).

Stratigraphic levels Cycle Species (specimen number) Sediment type Specimen size (mm) height × width Number of aliquots
(number of whorls)

V-3a 5 Viviparus (27/25,297) Coquina 28.1 × 16.2 (5) 24

V-4a 5 Viviparus (29/25,296) Molluscan sand 14.2 × 9.6 (5) 3
V-5a 5 Viviparus (10/7,020) Coquina 30.6 × 16.8 (5) 24
V-6a 4 Bellamya (26/43) Black mud Height 35 (5) 43
II-2a 4 Bellamya (1/1,909) Coquina 28 × 17 (5) 10
II-4 3 Viviparus (15/2,070) Coquina Height 27 24
II-5a 3 Viviparus (15/7,016) Black mud 40 × 25 (5) 13
II-5 and II-6 contacta 3 Viviparus (16/2,244) Black mud/ Coquina 14.6 × 9.8 (6) 7
II-6 L7a 3 Viviparus (14/2,051) Fluvial sediments Height 18 18
II-14 (MBB)a 2 Viviparus (28/25,300) Coquina Broken piece of 20 mm 3
a
Archaeological horizons with hominin artifacts (Goren-Inbar et al., 2000).
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B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44 37

Fig. 4. a–h: δ18O and δ13C data for individual Viviparid shells for time-series samples, plotted from the aperture (most recent) towards the apex (oldest), in the form of time
series of δ18O and δ13C values (a) and δ18O and δ13C cross plot (b). a: bed II-6 L7, b: contact between bed II-5 and bed II-6, c: bed II-5, d: bed II-4, e: bed II-2, f: bed V-6, g: bed V-5,
and h: bed V-3.
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38 B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44

Fig. 4 (continued).
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B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44
types including coquina, sand (molluscan shell sand), black mud, and
fluvial sediments from the sedimentary cycles 2–5 of Feibel (2004),
(Table 1).
5.2. Profiles of δ18O and δ13C isotope composition in individual shells
from GBY
The δ18O and δ13C profiles and δ18O vs δ13C cross plots of aliquots of
the individual shells are given in Fig. 4a–h (oldest 4a, youngest 4 h), for
8 of the samples with 7–43 aliquots. Samples of layers II-14 and V-4
yielded only three aliquots. They are not represented in the plots but
were included in the overall compilation (Fig. 5).
The profiles of the individual shells differ in both δ18O and δ13C.
Overall it is possible to identify three general types of profiles: 1)
relatively flat profiles for both δ18O and δ13C are present for: contact
between beds II-5 and II-6, (Fig. 4b); bed II-5 (Fig. 4c); bed II-2 (Fig. 4e).
2) Relatively flat profiles for δ18O with more pronounced variation in
δ13C for bed V-5 (Fig. 4g), and bed V-6 (Fig. 4f). 3) Moderate variations in
δ18O and stronger variations in δ13C for bed II-6 L7 (Fig. 4a), bed II-4
(Fig. 4d) and bed V-3 (Fig. 4h).
In all cases the youngest parts of the shell show relatively constant
values of both δ18O and δ13C except for sample II-4 (Fig. 4d) where the
profiles show increase in values of both δ18O and δ13C. Among the long
profiles, a synchronous reversal of the trends of δ18O and δ13C occurred
at least twice in profiles of layers V-5 (Fig. 4g), V-6 (Fig. 4f), II-4 (Fig. 4d)
and II-6 L7 (Fig. 4b).
5.3. The range of δ18O and δ13C values along the GBY sequence
The δ18O and δ13C of all shells investigated are presented in Fig. 5.
The overall range of values is from −10.5‰ to − 5.2‰ (VPDB) for
δ18O and from −9.2‰ to −4.8‰ (VPDB) for δ13C. Among the long
profiles the most scattered data for both δ18O and δ13C are from layer
V-6 (black mud) and the least variable is from bed II-4 (coquina),
belonging to cycles 4 and 3, respectively. The average values of δ18O of
the shell samples arranged through the GBY stratigraphy show a large
decrease towards the middle of cycle 3 and smaller fluctuations in
cycles 4 and 5, (Fig. 6). In most specimens the variability in δ18O is
small with standard deviation values between 0.2, mostly around 0.3
Fig. 5. Composite δ18O and δ13C of all samples investigated. Note the relatively low δ13C values of specimen at the contact II-5/6 V-6 and the relatively low values of δ18O of specimens
of beds II-2 and II-5.
39
and up to 0.8. The standard deviation of δ13C ranges between 0.2 and
1.4. However, the profiles have different patterns in various parts of
the shell. In one shell (bed V-6) (Fig. 4f), there are distinct differences
between early and late sections. Covariation of δ18O and δ13C values is
discernible in the sample from bed II-6 L7 (Fig. 4a) and some parts of
individual shells, but overall this is rare. The maxima and minima of
δ13C and δ18O in the time-series profiles occasionally coincide but
normally not. It seems therefore that the variations in δ18O and δ13C
are controlled by separate factors.
5.4. δ18O and δ13C patterns and values in relation to sediment type
The isotope profiles of sampled specimens are from several sediment
types (coquina, molluscan shell sand and black mud), (Table 1). Samples
from black mud show a specific pattern with more negative δ13C values
(Fig. 5). Samples of molluscan sand V-4, coquina II-2 and black mud II-5
have lower δ18O than the others. Therefore it seems that the relation
between the sediment characteristics and the isotope composition is
prominent for shells from black mud, these having more negative δ13C
values. Samples from the other sediment types do not show specific
isotopic characteristics.
6. Discussion
6.1. Occurrence of Viviparidae along the stratigraphic
beds — climatic implications
In our study no specimens of Viviparidae were detected in the
sedimentary sequence of GBY below the Matuyama–Brunhes Bound-
ary (MBB). The presence of other aquatic molluscs (Ashkenazi and
Mienis, 2005), ostracods (Rosenfeld et al., 2004) and aquatic
vegetation (Melamed, 1997, 2003) within the sedimentary sequence
below the MBB at GBY indicates that several layers were probably
deposited on the lake shore or swamps with rich vegetation, habitats
that seem to be suitable for Viviparidae. If there were taphonomic
effects such as of extreme acidity that dissolved the shells of the
Viviparidae (Ashkenazi and Mienis, 2005), then these would have
affected the other mollusc species and the ostracods as well. It seems
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40 B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44

Fig. 6. Composite stratigraphy at GBY (based on Fig. 11 in Feibel, 2004), with average values of δ18O and δ13C of individual Viviparus apameae galileae and Bellamya sp. from the
marked beds, and their content of organic carbon. The cycles (1–6) and the MBB are marked (R — reversed, N — normal). The main climate trends indicated are based on botanical
and zoological evidence.

therefore that other factors such as cold temperature, desiccation or
high salinity caused the absence of Viviparidae below the MBB.
Viviparus occur for the first time in the sequence of GBY after the
MBB, in layer II-14 in the first stages of cycle 2 (Fig. 6). The occurrence
of Bellamya sp. of either African or Asian origin (Prashad, 1928;
Brown, 1980) from bed II-7 towards the end of cycle 2, suggests that
the climate in this region was warmer than for previous layers,
enabling penetration of biological elements from Africa and/or Asia.
Bellamya coexists with Viviparus in the younger layers of the GBY
sequence. The genus Bellamya occur in the Lower Pleistocene (ca 1.7–
2.0 Ma) lacustrine sediments of Erq el-Ahmar (Sivan et al., 2006) but
not in the site of ‘Ubeidiya’ (ca 1.4 Ma, Tchernov, 1973, 1975) both in
the Lower Jordan Valley. The Pleistocene Hula (GBY Lake) and Erq el-
Ahmar lakes were freshwater lakes (~ 300 mg/l), while the salinity of
the water source of Ubeidiya was about 1000–1300 mg/l (Rosenthal
et al., 1989). The high salinity at Ubeidiya could account for the
absence of Viviparidae at that site. The appearance of Viviparidae
towards the end of cycle 2, after the MBB, could be the result of change
in climatic conditions and available habitats.
6.2. Intra- and inter cycle habitat change and climate variation
Our assessment of the climatic fluctuations through the six sedi-
mentary cycles and the individual beds within them (as described by
Feibel, 2004) is based on findings of plant and mollusc species
indicative of habitats and their respective ecology and provenance.
The first cycle starts with a basal fluvial conglomerate that is overlain
by coquina and later by gray mud. The bounding fluvial conglomerate at
the base of the sequence in cycle 1 is interpreted as reflecting a cool and
dry climate (Feibel, 2004). This is reinforced by the presence of the
elements of Palearctic and Euro–Sibirian origin in these sediments. The
oldest examined sample in the sequence is from the contact between
two beds, gravel and coquina (IV-24 and IV-25), which consists of a rich
assemblage of molluscs including several species of the genera Valvata,
Acroloxus and Oxyloma of Palearctic origin, that live in shallow water in
association with aquatic macrophytes. This sample contains several
ostracod species, parts of young crabs and small fish, which indicate a
shallow water and riparian habitat. Among the plants, oospores of Chara
sp., macrosporangiums of the aquatic fern Salvinia natans (L.) All. and,
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B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44
fruits of Trapa natans L. (both of Euro–Sibirian origin) and nutlets of two
Scirpus sp. together with remains of some riparian and dry-land plants
as Adonis sp., Ranunculus cf. marginatu D'Urv. and Vitis sylvestris C.C.
Gmel., indicate a shallow water or shoreline habitat. Although the flora
and fauna indicate a shallow water habitat suitable for Viviparidae, no
Viviparidae were detected in this assemblage.
The overlying bed III-11 contains a rich assemblage of plants,
molluscs, fish and crab remains indicative of a shallow productive
lakeside habitat. The presence of the tropical fern Azolla (Melamed,
2003) indicates at least a certain period of warm climate following the
cold/dry climate that prevailed during the sedimentation of the basal
conglomerate (Feibel, 2004). The successive beds show certain
fluctuations in specimen abundance with a trend of gradual decline
in aquatic mollusc species and the appearance of those typical of
shallow water, swamp and terrestrial habitats. These layers contain
relatively few plants, in comparison with the underlying bed (III-11).
They show a systematic decline in submerged plants and the presence
of emerging water plants such as Scirpus sp., Typha sp, Cladium
mariscus (L.) Pohl, and riparian plants like vine Vitis sylvestris
(Melamed, 2003). The emerging plants are resistant during periods
of desiccation, although their main biomass is above the water, their
root system still exploits water from the sediment. These beds include
two distinct palaeosols (beds III-6 and III-2). The beds between the
palaeosols (III-3 and III-4) contain only a few specimens that belong to
a very few mollusc species.
The overlying cycle 2, which contains the MBB was a moderate
transgressive phase. It starts with a drying wetland, accumulation of
shallow water submerged plant Naj as delilei Rouy (bed II-17) and
lignite (bed II-16). The lignite bed is devoid of mollusc shells or plant
seeds. These sediments deposited prior to the MBB represent relative
lacustrine lowstands (Feibel, 2004), conditions that are unsuitable for
Viviparidae. Bed II-14, which is the first bed above the MBB, contains a
rich assemblage of molluscs including the first appearance of Vivi-
parus. The younger layer II-11 and the subsequent sediments contain
abundant remains of the water fern Azolla, indicating a warm climate
(Melamed, 2003). The African or Asian gastropod species Bellamya
occurs for the first time at the end of this cycle in bed II-7. Its
appearance is accompanied by the occurrence of seeds of warm
Mediterranean plants such as Olea europaea and Ficus carica (Goren-
Inbar et al., 2002a) and the disappearance of the cold climate aquatic
plants Polygonum lap athifolium L. (Euro–Sibirian, Irano Turanian with
the most southern distribution range in the Mediterranean) and the
extinct Stratiotes intermedius (Harts) Chandler found in higher latitude
sites in Euro-Asia (Melamed, 2003). Moreover, among the tree
assemblages of GBY, Peryploca aphylla Decne and Ziziphus sp., both of
Sudanese origin, occur only during the warm period of bed II-6 (Goren-
Inbar et al., 2002a). Overall, the ratio of aquatic to dry-land vegetation
in GBY beds decreases from bed II-11 to the younger bed II-7
(Melamed, 1997). The warmer climate of beds II-7 and II-6 is reflected
in the increasing amount of seeds of the riparian plant Rubus sp., the
Mediterranean warm climate tree Olea europaea and the tropical plant
Cyperus articulatus L. (Melamed, 2003).
Cycles 3–5 represent oscillations of dry and wet climate (Feibel,
2004). Cycle 3 starts with layer (II-6) of fluvial conglomerate that
contains a diverse mollusc assemblage. As the lake desiccates towards
the end of the cycle, beds II-4 and II-3 contain very few mollusc species
including some land snails. The mollusc Melanoides tuberculata either of
north and east African or southern Asian origin, occurs in the site only in
bed IV-7 at the beginning of cycle 3 and exists at least until the end of
cycle 4 in bed V-6, indicating prevailing warm conditions. This species is
typically found in warm temperate to tropical freshwater bodies
(Livshits and Fishelson, 1983). Another example that reinforces the
evidence of a climate change in GBY from colder to warmer around the
MBB is the appearance of the bivalve Corbicula fluminalis (Müller, 1774)
an indicator of a temperate-warm climate (Meijer, 1985) and drying
saline habitat (Mania, 1995). At GBY it appears only in the younger beds
41
after the MBB, from layer II-6; the first layer of cycle 3. The biogeographic
distribution of ostracods is very similar to that of molluscs. At GBY the
distribution of 11 species of ostracods and 30 taxa of diatoms also
demonstrate changes from a cold climate with freshwater during
cycle 1, through a gradually warming phase, to a warmer climate
possibly with a small increase in water salinity in subsequent cycles
(Rosenfeld et al., 2004). This is corroborated by the negative trend of
δ18O in the ostracod Candona neglecta Sars, 1887 that is abundant in the
GBY samples (Rosenfeld et al., 2004).
Cycle 4 starts with bed II-2 that has the highest abundance of mollusc
species (33 species) indicating the most diverse habitat among the
examined layers. The last layer in this cycle (V-6), records the
desiccation phase of the lake marked by rich vegetation assemblage
(65 species), with diverse aquatic, swamp and shoreline vegetation
(Melamed, 2003). It also contains diverse mollusc species, animal bones
and hominin artifacts. The richness of species of this cycle is attributed to
desiccation that created numerous micro-habitats during the drying and
exposure of the shallow water and shore habitats. The desiccation of the
habitat in cycle 4 enabled hominin activity and artifact accumulation at
the site.
Cycle 5 starts with the coquina of bed V-5, accumulating a rich
mollusc assemblage including large quantities of Viviparus and Bellamya
species, animal bones and hominin artifacts. The living floor with
hominid activity together with molluscan shells accumulations, indicate
shallow water and shore environments. The bounding conglomerate of
cycle 6 consists of large basalt boulders correlated to the cool and dry OIS
18.2 (Feibel, 2004).
In summary, the combined flora, molluscs and ostracods assem-
blages in the sediments at GBY follow the five cycles recognized by their
sedimentological characteristics. (Feibel, 2004). The oldest beds in cycle
1 indicate a cold and dry climate with elements of Palearctic and
Siberian origin, with increasing desiccation towards the end of the cycle,
shown by the generation of several dry beds and two paleosol layers.
The warm climate continues into cycle 2 with a rise in the lake water
stand creating shallow water habitats with increasing abundance of
elements of swamp but also dry beds around the MBB. A further rise in
lake water stand and warming towards the end of the cycle and the
beginning of cycle 3 is indicated by the occurrence of Viviparids that
require deeper water, and the appearance of elements of African and
Sudanese elements. Cycles 4 and 5 show gentle fluctuations in the lake
water stand and the occurrence in close proximity of shallow and
deeper water elements including Viviparidae.
6.3. Profiles of the δ18O and δ13C in individual shells
Constant values and smooth patterns of both δ18O and δ13C were
observed in smooth parts of the shell; without evidence for a hiatus a
continuous shell growth of these parts is assumed. However, thicker
growth lines in various parts of the shell were interpreted as re-
presenting cessation of deposition, “winter rings” as suggested by van
der Spoel (1958). Variations in δ13C indicate variation in contribution
of organic derived and inorganic sources to the pool of DIC in the
water. The changes in δ13C reflect the productivity/decay variation
while the contribution of respired (metabolic) carbon seems to be
minor as discussed above.
On the basis of the systematic study of shells of recent Viviparus and
other freshwater gastropods in diverse environments (Schmitz and
Andreasson, 2001), the relative constancy of δ18O in the individual shells
can be taken as indication of low variability in the δ18O value of the
water and its temperature throughout the depositional history of the
shell or its examined portion. Some of our profiles differ, however, from
those of Schmitz and Andreasson (2001) which are marked by
fluctuations of between 2 and 5‰ in δ18O. The rapid large fluctuations
of δ18O in the specimen from bed V-6 (Fig. 4f) in the part marked by a
smooth 4‰ variation in δ13C is difficult to explain. Equally pronounced
excursions in δ13C were reported from Viviparus from Johns River
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42 B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44
Florida and Bei Kenya (Schmitz and Andreasson, 2001). However, the
accompanying δ18O profiles are marked by fluctuations too, which
suggest a pronounced kinetic effect.
6.4. Ontogenetic and ecological factors relevant to interpretation of
isotopic results
Viviparidae give birth to fully developed young with shell (Browne,
1978). In the extant Viviparus georgianus (Lea, 1834) embryos develop
over 9 months and the fully formed young with three whorls are
released to water during April–May (Browne, 1978; Jokinen et al.,
1982). Hatched young Viviparid gastropods tend to grow among
submerged vegetation and algae. In the Hula Valley this environment
is stable during April–May until temperatures start to rise in June–
August (Ashkenazi et al., 1999).
We interpret the first drop in δ13C values in the profiles of specimens
from beds V-3, V-5, V-6, II-4 and II-6 L7 having a fluctuating pattern, as
an increase in the contribution of carbon of organic origin. This happens
at the end of summer when the submerged macrophytes cease to grow
and decompose providing ample amounts of litter, which may also
increase the metabolic rate of the Viviparidae that consumes it. The
pattern of relatively moderate fluctuations of δ18O is attributed to a
thermal stability and consistency of the δ18O values of the lake water.
These features indicate essentially a constant inflow and consistent
isotopic composition, in analogy to the subsequent Hula Lake. The
section of the shell between consecutive maxima or minima in δ13C,
such as seen in the specimens from beds V-5, V-6, II-4, II-5 and II-6 L7
represents a year's growth. The latest portions of the profiles in these
specimens are marked by increase in both δ18O and δ13C (Fig. 4c)
suggesting the effect of evaporation although biological controls cannot
be ruled out.
The profiles of the Viviparid specimens from beds V-5, V-6, II-4
and II-6 L7 are interpreted as following the present day environ-
mental cycle in shallow lakes in the Hula Valley (Ashkenazi et al.,
1999: Figs. 4 and 7). We assume that the parallel phase of δ18O and
δ13C of young Viviparidae growth profile included the period spring–
autumn when the released young could encounter stable, vegeta-
tion-rich aquatic habitat, until autumn, when the shell δ18O values
increase and δ13C decrease. The water temperature of the shallow
lake in the Hula Valley drops in winter to 10–12 °C limiting the
growth of aquatic plant until the next spring (Ashkenazi et al., 1999).
The aquatic plant decay process of the following year results in the
continuation of the drop in δ13C and a slight increase in δ18O.
Although the studied Viviparid shells show the thicker lines of
cessation of growth these lines are not expressed in the isotope
profiles, a phenomenon also observed by Schmitz and Andreasson
(2001). The variability in the seasonally changing patterns of δ13C
and δ18O in Viviparid shells are possibly effects of micro-habitats.
Such an effect of micro-habitat is seen in δ18O and δ13C values of
Ostracods and the Bivalve Pisidium spp. from the Ammersee and
Starenberger See in Southern Germany investigated for the season-
ality features of their growth as different species grow during
different seasons (Von Grafenstein et al., 1999). However, it is
difficult to test whether this is the case for the viviparids at GBY.
Szymanek (2005) showed that the fossil Viviparus diluvianus (Kunth,
1865) from the same site in Poland formed at different periods show
differences in size. Shells associated with cold periods are less
abundant, have smaller mean height and are less slender then those
from beds deposited under conditions of more oceanic climate. The
interpretation of the seasonality pattern in the shells with long time
series can provide an estimate of yearly growth rate. In the case of
the specimen from layer II-5 the intervals between two extreme
values of δ18O and δ13C are 5–10 aliquots are interpreted as growth of
ca 15–30 mm/year. This is in agreement with the extant Viviparus
georgianus annual growth rate of 10–30 mm (Browne, 1978).
6.5. The range and values of δ18O and δ13C
The overall range of δ18O is ca −10.5 to −5.2‰, and of δ13C ca −9.2
to −4.8‰ (VPDB). The spread of data of δ18O in eight out of the ten
samples is narrow, with a mean value around −7.5‰ (VPDB) with two
samples (II-5 black mud and II-2 molluscan sand) being more negative,
around −9.2 and −8.5‰ (VPDB) respectively. Two samples II-5 (black
mud) and the contact between II-5 and II-6 (black mud and coquina)
have δ13C values lower than the rest.
The samples from black mud horizons have lower δ13C values
compared to other samples, in line with the organic carbon content in
the sediments given in Fig. 6. The difference in δ18O values indicates
that deposition of organic rich sediments may have taken place under
different hydrological regimes, which may reflect climatic conditions.
Sample II-5 shows the most negative average δ18O value of − 9.1‰,
whilst the underlying contact between II-5 and II-6 has a less negative
value of δ18O of − 6.9‰, which is similar to that of the Viviparidae
samples from some coquina beds. The position of this sample in the
contact between coquina and black mud is thus reflected in the
isotopic features. It seems therefore that the average δ13C values of the
shells reflect the average contribution of organic derived carbon and
hence characterizes the local habitat of the gastropod, while the time
series reflect the seasonal variability of the relative contribution from
organic source (plant litter) and hence the seasonal changes occurring
in the habitat with the additional change in metabolic rate. Two
samples have particularly low δ13C values II-5, and contact II5/II6, but
the other samples have the same range of composition. This indicates
that the hydrological and climatic conditions as represented by these
samples are relatively stable with the intra-shell variations in the
range of the variation among the averages of the different shells.
The δ18O values of the ostracod Candona neglecta from cycles 2 to
5 at GBY (Rosenfeld et al., 2004) have a narrow range of − 5.3 to
− 3.8‰ (VPDB) with the more negative values at the lower part and
the top of the section. Samples were taken from grey marls and
coquinas in cycle 1, lower part of cycle 2 and black mud in cycles 2 to
5. The range of δ13C is much wider − 8.6 to + 3.6‰ with overall lower
values in cycles 2 to 4 than those from the base and top of the section.
This variability is interesting in view of the similarity of the sediment
(black mud) and the fact that ostracods form their shell during a very
short period in the same season.
The variable nature of the sediment indicates the changes in water
level and productivity of the lake and hence the controlling
hydrological regime, which seem quite pronounced. The comparison
between the average δ18O values of shells for the sedimentary cycles
reveals a large interval in cycle 3, a smaller one in cycle 4 an even
smaller one in cycle 5. Overall the correlation between the
stratigraphy of the sedimentary beds and the synthetic curve of the
OIS given by Feibel (2004) is corroborated by the present results with
the large isotopic differences in the lower part cycles 2 and 3 (2.2‰)
and diminish within cycles 4 and 5 (1.2‰) and are even lower in cycle
5. These agree with the model of Bassinot et al. (1994) with the
biggest difference between OIS 19.1 and 18.4 of ca 2‰, diminishing
upwards (see Fig. 6) The model based on oceanwater δ18O represents
therefore a change in the source of the rainwater that eventually
precipitates in the catchment, and thus the changes recorded at GBY
can be related to the global climate system.
The overall relationship between isotope composition and environ-
mental conditions, as reflected in the types of host sediment, was
discussed above. However the details of the δ18O and δ13C characteristics
recorded in the Viviparidae at GBY above the MBB can be interpreted
with reference to those recorded in extant molluscs including Vivipar-
idae originating from various zoogeographic zones reported by Schmitz
and Andreasson (2001). On the basis of this comparison we suggest that
the climate prevailing in GBY during the deposition of the sedimentary
beds following MBB was oscillating between temperate, subtropical to
humid tropical. In the wider context of the variety of the habitats the
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B. Spiro et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 282 (2009) 32–44
biogeographic origin of floral and molluscan elements agrees in broad
terms with the trends in isotope composition of the Viviparidae from
these climatic zones. More pronounced differences among the biological
assemblages which occupy the shore belt environment seem to reflect a
higher sensitivity to changes occurring in this zone. In the more off-shore
sediments, at up to 3 m water depth, which are the habitat of Viviparidae,
these changes are less pronounced. While the overall trend is from cooler
to warmer climate across the MBB, the fluctuations recorded by the
distribution of biological elements and the isotope values of Viviparidae
diminish through the GBY sedimentary cycles. Overall the correlation
between the stratigraphy of the sedimentary beds and the synthetic
curve of the OIS given by Feibel (2004) is corroborated by the present
results with the large isotopic differences the lower part, namely cycles 2
and 3 (2.2‰) diminishing within cycles 4 and 5 (up to 1.2‰) in
agreement with the model of Bassinot et al. (1994). This model is of
oceanwater δ18O variation in relation to ice volume and therefore
applicable to the δ18O of the rain water.
Based on the isotopic composition of Viviparid molluscs and the
occurrence of indicative plant and molluscan elements in the GBY
beds, we suggest here that the geophysical-magnetic phenomenon
MBB coincides with a significant environmental change. The prevail-
ing climate fluctuated from cold and dry in the oldest beds of the site
changing to warmer and more humid fluctuating climate following
the MBB. This relationship may indicate a link between both warming
of climate and decrease of salinity in the GBY lake and an increase in
hominin activity.
7. Summary
The sedimentary succession at Gesher Benot Ya'aqov (GBY) provides
an environmental record of a lakeside habitat over ca 100 Ka crossing
the Matuyama–Brunhes Boundary (MBB) at 780 Ka. This boundary
occurs in the second sedimentary cycle of a series of six, which is
bounded by fluvial conglomerates at the bottom and top of the
succession. Each cycle starts with lake-shore sediments followed by
deeper water ones. The flora, mollusc and ostracod assemblages occur
along the entire sequence of GBY while the viviparids studied here occur
only above the MBB in cycle 2. The fauna and flora show a change in
species composition, richness, abundance, biogeographic origin and
habitat requirement. These changes indicate a trend of climate warming
at the level of the MBB and more moderate changes within the
subsequent cycles revealing a link between type of sediment and its
floral and mollusc record. The near absence of Viviparidae below this
boundary is attributed to cooler climate at the base of the section
characterized by the presence of Euro–Sibirian and Palearctic elements
and desiccation (paleo sols) in the sediments of cycle 1. The appearance
of the Viviparidae of African and/or Asian origin in cycle 2 marks the
increase in water level and temperatures reflecting a warm and humid
climate, following the MBB at the end of a prolonged dry climate. These
observations are the basis for the new suggestion that the MBB which is
an important geophysical-magnetic phenomenon coincides with a
significant environmental change in this region.
The δ18O and δ13C records provide a different insight into the
palaeoenvironment and its change. The time series of δ18O and δ13C in
shells of Viviparidae, Viviparus apameae galileae and Bellamya with
relatively large shells and longevity, provide a record of seasonal and
annual fluctuations in environmental conditions. The within-shell
range of δ13C values (maximal range −9.2 to −4.8‰ VPDB) reflects a
large variability in the contribution of organic derived carbon with
possible slight influence by metabolic carbon. Specimens from black
mud beds rich in organic matter have δ13C values in the lower part of
this range reflecting the enhanced contribution from this source. The
overall range of average δ18O values for the individual shells is −9.1 to
−6.9‰ VPDB with the lowest values from black mud bed. However,
there does not seem to be a clear correlation between the type of
sediment and the δ18O values of the viviparids shells.
43
The small variability of intra-shell δ18O values indicates near
constant conditions within the water body on a seasonal and inter-
annual scale, which suggests that the lake is fed by constant source
(river or springs) with consistent δ18O, not by floods. The cyclic
pattern of the δ13C values probably corresponds to the seasonal
growth and decay of aquatic macrophytes in spring and autumn
respectively, the season of death of the individual gastropods and
their rate of shell growth of 15–30 mm/year. Moreover, the lack of
within-shell covariation of δ18O and δ13C suggests that evaporation
was not a considerable factor and that the isotope systematics is a
good record of the prevailing environmental conditions.
The record of shell — average δ18O values and their respective
stratigraphic positions corroborates the interpretation of the sedi-
ment cycles as representing change in climate and their correlation
with the OIS stages 20 to 18 as proposed by Feibel (2004). The
recorded fluctuations show the same trends as the model of Bassinot
et al. (1994) for variation in δ18O ocean waters with a decrease in
intensity towards the later cycles. This agreement of trends of δ18O of
ocean waters and viviparids from the lacustrine sediments of GBY
supports the notion that the δ18O of the viviparids provide a good
representation of the hydrology of a catchment dictated by the
atmospheric precipitation with a link to global climate record for that
time interval. The major change occurs in cycle 2, a transition from
cold dry to increasingly warm and fluctuating dry/humid climate.
These observations are the basis for the suggestion that the MBB
coincides with a significant environmental change in this region.
The combined information derived from the study of plants and
stable isotope analysis of viviparidae in a single site where they occur
over ca 60 Ka suggests that the δ18O record agrees with the record of
habitats and provides an environmental record of the lacustrine
hydrology and climate, while the δ13C record accentuates the annual
productivity cycle of the lake, both contributing to the palaeoenvir-
onmental reconstruction.
Acknowledgements
Prof. Naama Goren-Inbar from the Institute of Archaeology, the
Hebrew University, is thanked for the sample material and information
from the GBY site, that made the present study possible. Thanks are due
to Gabi Laron from the same institute for the photographs. Thanks are
expressed to Adi Ben-Nun from the GIS unit, Department of Evolution,
Systematics and Ecology, the Hebrew University, Jerusalem for the
preparation of the map and to Dr Neta Spiro for the preparation of the
graphics. The stable isotope analyses were carried out at the stable
isotope laboratory, University College London and thanks are due to
Prof. T. Atkinson. Pilot studies were supported by the RING Foundation
2003–2004 and by the Irene Levi Sala CARE Archaeological Foundation,
2005–2006. The final part of this study was carried out by an ongoing
grant awarded by the Israel Science Foundation (grant no. 300/06) to
the Center of Excellence — project title: “The effect of Climate Change on
the Environment and Hominins of the Upper Jordan Valley between ca.
800 Ka and 700 Ka ago as a Basis for Prediction of Future Scenarios.”
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