Jrowland Final

INTER- AND INTRA-TOOTH ISOTOPIC VARIATION IN
MAMMALIAN TOOTH ENAMEL FROM WESTERN ISRAEL:

IMPLICATIONS FOR PALEOENVIRONMENTAL AND
PALEOCLIMATE CHANGE OVER THE PAST 350 KYR
by
Jessica C. Rowland
A Prepublication Manuscript Submitted to the Faculty of the
DEPARTMENT OF GEOSCIENCES
In Partial Fulfillment of the Requirements

for the Degree of
MASTER OF SCIENCE
In the Graduate College

THE UNIVERSITY OF ARIZONA
2006
STATEMENT BY THE AUTHOR
This manuscript, prepared for publication in the Journal of Archaeological Science, has
been submitted in partial fulfillment of requirements for the Master of Science degree at
The University of Arizona. A copy of the manuscript is filed in the Antevs Reading
Room to be made available to borrowers.
Brief quotations from this manuscript are allowable without special permission, provided
that accurate acknowledgment of the source is made. Requests for permission for
extended quotation from or reproduction of this manuscript in whole or in part may be
granted by the Department of Geosciences when the proposed use of the material is in the
interests of scholarship. In all other instances, however, permission must be obtained
from the author.
_______________________________________________________ _____________
Jessica C. Rowland
APPROVAL BY RESEARCH COMMITTEE
As members of the Research Committee, we recommend that this prepublication

manuscript be accepted as fulfilling the research requirement for the degree of Master of
Science.
_______________________________________________________ _____________
Jay Quade, Major Advisor
_______________________________________________________ _____________
Mary C. Stiner
_______________________________________________________ _____________
David L. Dettman
_______________________________________________________ _____________
Vance T. Holliday
2
Abstract
Stable isotope values (δ13C, δ18O) of large herbivore bioapatite are used to
examine paleoenvironmental and paleoclimate change in the coastal hills of the Levant
region. We draw upon archaeological faunal records of fallow deer (Dama) and mountain
gazelle (Gazella) from Qesem Cave, Hayonim Cave and Meged Rockshelter (Israel) that
span the past ~350 kyr. Because faunal records indicate that a sudden dispersal of
Gazella from the Afro-Arabian biotic province into the Levant region occurred around
200 ka BP, we investigate the possibility that the proportion of the two species was
modulated over time by climate and/or environmental change. δ13CPDB values of fossil
molar enamel from all time periods fall between -14.2 and -7.3‰, indicating that large
herbivore diet was dominated by C3 plants. Inter-tooth δ13C data reveal that Dama and
Gazella occupied discrete niches that remained relatively stable over time, despite
increasingly arid conditions and glacial-interglacial climate fluctuations. Enamel
δ18OSMOW values from the same time series have a range of +26.2 to +34.9‰, with
average δ18OSMOW values gradually increasing over time by ~4‰. Taken together, these
results imply that climate change likely was not the driving factor behind the rapid influx
of Gazella into the Levant region ~200 ka BP. A comparison of bioapatite δ18O values
with reconstructed meteoric water δ18O values yields an enrichment (εbioapatite-water) on the
order of 29-37‰, much larger than that predicted by recent physiologic models of δ18O
values. This finding points toward significant species-specific enrichment of δ18OSMOW
values, likely due to diet preferences and physiologic factors.
Introduction
The Levant is a biogeographic corridor between Africa and Eurasia that has
served as an important region for both northward and southward dispersals of hominids
and other biota since the early Neogene (Tchernov, 1992). There has long been much
interest in the area for its diverse hominin record (see Akazawa et al., 1992 and Akazawa
et al., 1998 for references therein), and related records of paleoenvironmental and
paleoclimate reconstruction. Previous studies have drawn upon evidence from lake
records (Bartov et al., 2002; Ginat et al., 2003; Hasse-Schramm et al., 2004; Hazan et al.,
2005), soil studies (Magaritz, 1986; Gvirtzman & Wieder, 2001; Frechen et al., 2004),
pollen records (Weinstein-Evron, 1987; Horowitz, 1989; Albert et al., 2003), speleothem
3
records (Bar-Matthews et al., 1999; Frumkin et al., 1999; 2000) and from Natufian and
Neolithic tooth assemblages (Shahack-Gross et al., 1999; Richards et al., 2003).
In this paper we examine a time series of sub-fossil ungulate teeth from Qesem
Cave, Hayonim Cave and Meged Rockshelter (Fig.1) that spans approximately the last
350 kyr. Stable carbon and oxygen isotope analyses of tooth enamel (bioapatite) have
been widely used to reconstruct ancient climate and environments (e.g., Ayliffe &
Chivas, 1990; Quade et al., 1992; Fricke et al., 1998a; Koch et al., 1998; Gadbury et al.,
2000), because enamel is highly resistant to diagenesis (Lee-Thorp & van der Merwe,
1991; Ayliffe et al., 1994; Shahack-Gross et al., 1999). Inter- and intra-tooth isotopic
change are investigated in this paper, in order to gain insight into both long-term climatic
change and short-term seasonal variation (e.g., Fricke and O’Neil, 1996; Kohn et al.,
1998; Zazzo et al., 2002; Balasse et al., 2003).
Considerable turnover has been noted in mammalian communities of the region
during the late Pleistocene. Bate (1937a, 1937b) called upon climate change to explain
the decline of fallow deer (Dama mesopotamica) and its replacement by mountain gazelle
(Gazella gazella) in the Near East, but this claim remains controversial. In our study area,
Dama remains dominate the Lower Paleolithic sequences, whereas Gazella become more
prevalent through the Middle and Upper Paleolithic layers. Zooarchaeological evidence
indicates that there was a rather sudden dispersal of Gazella from the Afro-Arabian biotic
province into the Mediterranean hills of the Levant region around 200 ka BP (Stiner,
2005; unpublished data 2006). The observed variation in the Mediterranean faunal series
could be potentially explained by 1) changes in human subsistence behavior and prey
selection, or by 2) climate-driven environmental changes. Here we use stable carbon and
4
oxygen isotope analyses to investigate if the proportion of Gazella with respect to Dama
in the Levant region is climatically modulated. In order to better understand the context
of the human evolutionary record in the region, it is critical to closely examine
mechanisms behind mammalian dispersals that were possibly concurrent with northward
movements of early modern human populations into the Levant.
Background
Site Locations, Chronologies, and Regional Climate
Qesem Cave is located ~12 km east of Tel Aviv, Israel, at ~90 m above sea level
(asl) (Fig. 1). The cave is formed in late Cretaceous limestone, and contains both
archaeological and natural deposits, including extensive calcite flowstones. Uranium
series (230Th/234U/238U) dating of a massive flowstone and a speleothem crust that
partially cover the lower and upper Acheulo-Yabrudian layers in the eastern section of
the cave has provided a tentative chronology of late Lower Paleolithic occupation (Barkai
et al., 2003). The massive flowstone, deposited from 382±37-207±12 ka BP, pre-dates
most of the Acheulo-Yabrudian layers. The flowstone is directly overlain by a thin
archaeological deposit, which accumulated during a hiatus in speleothem deposition from
207±12-152±3 ka BP.
The ~207 ka BP age estimate for the upper Acheulo-Yabrudian (Layer 2) is
generally accepted, but the correlation between the speleothem ages and the lower Qesem
stratigraphy (Layers 3 and 4) is best described as provisional. Barkai et al. (2003)
acknowledge that human occupation likely took place simultaneously with speleothem
deposition. Hence, based on the archaeological materials in Layers 3 and 4 and the dates
5
of other contemporaneous sites in the region (Bar-Yosef, 1998), we assume that the age
range for the lower Acheulo-Yabrudian deposits is ~300-350 ka BP. For the purposes of
this study, the mammalian teeth from Qesem Cave are divided into two approximate age
groups of 300-350 ka BP and 200-215 ka BP.
Hayonim Cave and Meged Rockshelter (Fig. 1) are located in the western Galilee
of Israel, approximately 30 km northeast of Haifa, at an elevation of ~250 m asl. The two
sites are about 1 km apart, and are formed in late Cretaceous limestone (Kuhn et al.,
2004). Hayonim Cave contains archaeological assemblages from the Middle Paleolithic
to Natufian periods, whereas Meged Rockshelter contains assemblages that represent the
terminal Upper Paleolithic and early Epipaleolithic periods. Middle Paleolithic
chronology at Hayonim has been determined by uranium-series dating of speleothems
and teeth (Rink et al., 2004), thermoluminescence dating of burnt flints (Valladas et al.,
1998), and electron spin resonance (ESR) dating of tooth enamel (Schwarcz & Rink,
1998; Rink et al., 2004). Upper Paleolithic and Epipaleolithic deposits at Hayonim have
been dated by radiocarbon methods (Bar-Yosef, 1991; Housley, 1994; Phillips, 1994).
Age estimates for Meged Rockshelter are based on AMS radiocarbon dates of charcoal,
and indicate that occupation centered on the Last Glacial Maximum (Kuhn et al., 2004).
All chronologies are presented in Table 1.
Qesem, Hayonim and Meged are situated within the Mediterranean climate belt, a
region characterized by warm, dry summers and mild, wet winters. Tooth samples were
selected specifically from these sites because they share similar ecogeographic settings.
Annual precipitation in the area in which the archaeological sites are located is about 550
mm, and falls mainly between November and March (IAEA/WMO, 2004). Average
6
January and July temperatures are 12°C and 25°C, respectively. Modern vegetation is
diverse, composed mainly of Mediterranean woodland (oak, carob and terebinth trees)
interspersed with stands of wild wheat and barley (Zohary, 1973).
Tooth Enamel Formation and Stable Isotope Analysis
Tooth enamel (bioapatite) is highly resistant to isotopic alteration compared to
dentine and bone (Lee-Thorp & van der Merwe, 1991; Ayliffe et al., 1994; Shahack-
Gross et al., 1999). It is composed of tightly packed crystallites of biogenic
hydroxyapatite (Ca10[PO4,CO3]6[OH]2) that are ~50-100 nm in diameter and >1000nm in
length (Hillson, 1996). Tooth enamel matrix initially has a high organic content, which is
mineralized gradually from the crown to the root of the tooth, and from the enamel-
dentine junction outward, during the process of amelogenesis (Fricke & O’Neil, 1996;
Kohn et al., 1998; Zazzo et al., 2005). A record of temporal isotopic change is preserved
along the growth axis of a mammalian tooth, representing a partially time-averaged
archive of ancient seasonality (Passey and Cerling, 2002; Passey et al., 2005). Intra-tooth
microsampling can provide a record of isotopic variation during the formation of the
tooth enamel (e.g., Fricke and O’Neil, 1996; Kohn et al., 1998; Zazzo et al., 2002;
Balasse et al., 2003).
The oxygen isotope composition of tooth enamel (or δ18Obioapatite, expressed in the
standard ‰ notation) is determined by the δ18O value of herbivore body water (δ18Obw),
which is primarily influenced by ingested drinking water and leaf water (Kohn et al.,
1996; Fricke et al., 1998b). For homeotherms (mammals that maintain constant body
7
temperature ~37°C), the measured fractionation factor (α) between enamel phosphate and
body water is represented by:
δ18OPO + 1000
α PO 4 −bw
= 18 4
(1)
δ Obw + 1000
and is equal to 1.0178 (Luz & Kolodny, 1985), whereas the estimated isotopic
fractionation factor between enamel carbonate and body water (αCO3-bw) is 1.026 (Bryant
et al., 1996b; Iacumin et al., 1996). In many studies, δ18Obw is assumed to be to the same
as the δ18O value of meteoric water (δ18Omw). However, species-specific diets and
physiological effects may influence the oxygen isotope enrichment factor between
ingested water and enamel carbonate (αCO3-water) (Ayliffe & Chivas, 1990; Bryant &
Froelich, 1995; Kohn, 1996; Kohn et al., 1996; 1998; Fricke et al., 1998b), leading to
αCO3-water > 1.026. Herbivores that require a regular intake of surface water and consume
mainly plant stems will likely have δ18Obioapatite values that are close to local δ18Omw
values, whereas drought-tolerant herbivores that consume evaporated waters and plant
leaves will have δ18Obioapatite values that increase with aridity (Levin et al., 2006). Leaf
water (δ18Olw) can be evaporatively enriched in 18O by 10-25‰ in comparison to stem
and local meteoric water (Dongmann et al., 1974).
The carbon isotope composition of mammalian tooth enamel (δ13Cbioapatite) is
correlated with the type of vegetation that an ungulate consumes (Lee-Thorp & van der
Merwe, 1987). C4 plants, such as warm-season or tropical grasses and some shrubs, have
δ13C values (δ13Cvegetation) around -10 to -14‰. C3 plants, which include trees, most
shrubs, and cool-season grasses, tend to have δ13C values ranging from -21 to -32‰. δ13C
values of C3 plants in arid environments are usually slightly higher than those living
8
under less water-stressed conditions (Ehleringer et al., 1992). Because precipitation in the
Mediterranean region occurs mainly during the winter, cool-season C3 plants are the
dominant type of vegetation today (Shomer-Ilan et al., 1981), as they were in the past
(Peyron et al., 1998; Frumkin et al., 2000; Guiot et al., 2000). As a result, little change
through time in δ13Cbioapatite values is to be expected, although minor variations may
indicate changes in aridity. For herbivorous ruminants, the measured carbon isotope
enrichment (εbioapatite-diet) between tooth enamel and diet is ~14 - 15‰ (Cerling and Harris,
1999; Passey et al., 2005).
Material
Fallow deer (Dama mesopotamica) and mountain gazelle (Gazella gazella)
molars were chosen for isotopic analysis. Third molars (M3) were preferred for this
study, although some third and fourth premolars (P3, P4) and second molars (M2) were
sampled when M3s were not available (Appendix 1). M3s and P4s are most suitable for
isotopic work because these teeth are late forming, and their enamel isotopic
compositions are not influenced by nursing and weaning (Bryant et al., 1996a; Kohn et
al., 1998). In order to obtain the longest possible temporal sequences, the highest
crowned (or, least worn) teeth were selected. Both upper and lower molars were sampled,
even though a slight offset in timing of growth of these teeth (Balasse et al., 2003) may
introduce some variability into our observations. Depending on the species analyzed, the
amount of time archived in a single M3 is on the order of a few months to one or two
seasonal cycles, (Fricke & O’Neil, 1996; Kohn et al., 1998; Balasse et al., 2003). This
issue will be discussed in more detail below.
9
Methods
Fifty-two ungulate teeth (558 total analyses) were prepared for oxygen (δ18O) and
carbon (δ13C) isotope measurements of the structural carbonate component of bioapatite
(Land et al., 1980). The enamel surface of each molar was first cleaned of adhering
sediment and organic matter using a stationary drill, and the highest loph was selected for
analysis. Microsampling was performed from the apex to cervix of the tooth, under a
binocular microscope with a 0.9 mm diamond-tipped drill bit. Furrows approximately 1
mm wide were drilled perpendicular to the growth-axis of the tooth at 2-3 mm intervals
(see Fricke & O’Neil, 1996 and Zazzo et al., 2002 for comparable microsampling
techniques). At this level of sampling, some attenuation or averaging of the total isotopic
amplitude present in the tooth is likely. Roughly 0.7 - 1 mg of enamel powder was
recovered from each furrow. Careful drilling and visual inspection of the enamel powder
ensured that dentine was excluded from the sample.
Powdered enamel is typically pretreated with dilute sodium hypochlorite (NaOCl)
or hydrogen peroxide (H2O2) to remove organic matter, followed by leaching with very
dilute acetic acid in order to dissolve secondary mineral contaminants (Koch et al., 1997),
leaving purified “structural carbonate” for analysis. The enamel microsamples used in
this study are so small (≤1 mg) that pretreatment often left insufficient sample for
analysis. This raised the question of the necessity of the pretreatments for the enamel.
The pretreatment procedure is certainly vital for analysis of bone (Lee-Thorp & van der
Merwe, 1991; Ayliffe et al., 1994), but unlike bone, enamel contains very little organic
matrix or little apparent non-structural carbonate (Hillson, 1986; Koch, 1998). To test the
effects of pretreatment on δ13Cbioapatite and δ18Obioapatite values, we carried out the following
10
experiments on four additional teeth of various ages from a range of geographic regions.
The fossil tooth samples, in order of increasing age, are from the following mammals:
Mammuthus columbi (~11-12 ka; Las Vegas, Nevada area); Equus caballus (~28 ka; Val
Boi, Portugal); and a Gomphothere species (~11 Ma; Pascalar, Turkey). A modern tooth
of a wild horse (Equus kiang) from the high Tibetan Plateau (near Lhasa, Tibet) was also
sampled. Enamel samples were cleaned and separated from dentine with a hand-held
dental drill, and fully homogenized by grinding in a mortar. The powdered bulk enamel
from each tooth was subsequently divided into four aliquots that were subjected either to
a) no pretreatment; b) 2% NaOCl for 12 hours; c) 2% NaOCl for 12 hours, followed by
0.1 M acetic acid for 2 hours; or d) 0.1 M acetic acid for 2 hours. All treated aliquots
were rinsed and centrifuged five times with distilled water, and dried at 50°C. Five to ten
samples of each aliquot were analyzed.
The δ13Cbioapatite and δ18Obioapatite values were measured using an automated
carbonate preparation device (KIEL-III) coupled to a gas-ratio mass spectrometer
(Finnigan MAT 252). Powdered samples were reacted with dehydrated phosphoric acid
under vacuum at 70°C. The isotope ratio measurement is calibrated based on repeated
measurements of NBS-19 and NBS-18, and precision is ±0.1‰ for δ18O and ±0.06‰ for
δ13C (1σ).
Vegetation samples (57 analyses) were collected from the vicinity of the three
archaeological sites from which the fossil teeth were recovered, and identifications follow
Zohary and Feinbrun-Dothan (1966-1986). Plant matter was pretreated with 2 M HCl,
rinsed with deionized water and dried at 50°C. Organic δ13C values were measured using
an automated CHN analyzer (Costech) coupled to a continuous-flow mass spectrometer
11
(Finnigan Delta Plus XL). Internal lab standards are calibrated relative to NBS-22 and
USGS-24, and precision of repeated internal standards is ±0.09‰ for δ13C (1σ).
In this paper, tooth enamel oxygen isotope results are presented using standard
δ‰ notation relative to VSMOW, whereas δ13C values of tooth enamel and organic
matter are reported relative to VPDB.
Results
Pretreatment experiment
Previous pretreatment experiments on tooth enamel (Lee-Thorpe & van der
Merwe, 1991; Koch et al., 1997) reveal that certain pretreatments modestly affect the
isotopic composition of enamel. Thus, it is critical to quantify the effect that pretreatment
procedures have on enamel of various ages, and to determine if pretreatment is necessary
at all. Comparison of the unpretreated (a) and fully pretreated (c) enamel aliquots
mentioned above is most relevant to this discussion (Figs. 2a and 2b).
In general, unpretreated fossil enamel has isotopic values that are slightly depleted
in 18O and enriched in 13C compared to fully pretreated fossil enamel. Although neither
the mean δ18Obioapatite nor δ13Cbioapatite values of the unpretreated and fully pretreated fossil
tooth enamel (Mammuthus columbi, Equus caballus, and Gomphothere species) are
statistically within error at the 95% confidence level, the differences between the mean
δ18Obioapatite and δ13Cbioapatite values are relatively small, ranging from 0.14-0.39‰ and
0.06-0.53‰, respectively (Table 2). The difference between the mean isotopic values of
the unpretreated and fully pretreated enamel samples is much less than the uncertainty
associated with other aspects of this study (e.g., estimating past δ18Omw values). The
12
modern Equus kiang tooth gives statistically indistinguishable (at the 95% confidence
level) mean δ18Obioapatite results for the unpretreated and fully pretreated aliquots, but the
δ13Cbioapatite values are 0.18‰ different. In conclusion, we found that the conventional
enamel pretreatment procedure is unnecessary in the context of our investigation (as in
Zazzo et al., 2005).
Temporal trends in isotopic values
Average δ13Cbioapatite values of Dama and Gazella fall into two distinct groups that
show relatively little change over time (Fig. 3a). Average δ13Cbioapatite values for Dama
fall in the range of -10 to -12‰, whereas average δ13Cbioapatite values for Gazella are
around -12 to -14‰. Generally, δ13Cbioapatite values of Gazella are lower and δ18Obioapatite
values higher than in Dama enamel. Average δ18Obioapatite values are much more variable
than the δ13Cbioapatite values, and range from +33.6 to +27.5‰ (Fig. 3b). Average
δ18Obioapatite values of both Dama and Gazella appear to increase gradually from ~350 to
20 ka BP, after which time they decrease.
Intra-tooth variation
The isotopic values of intra-tooth enamel microsamples vary substantially. The
δ13Cbioapatite values from all time periods fall between -14.2 and -7.3‰, and δ18Obioapatite
values from the same time series have a range of +26.2 to +34.9‰. The range of intra-
tooth δ13Cbioapatite and δ18Obioapatite values is variable through time, from 0.4-3.6‰ and 0.3-
4.1‰, respectively. Overall, intra-tooth isotopic variation in δ18Obioapatite values is greater
than that of δ 13Cbioapatite. Average intra-tooth variance over the entire time series is 2.3‰
13
for the δ18Obioapatite values, and 1.2‰ for the δ13Cbioapatite values. The cyclic variation in
δ18Obioapatite values likely reflects the seasonal cycle, with the highest δ18Obioapatite values
during the summer months and lowest during the winter months. The maximum
“seasonal” range of δ18Obioapatite values (4.1‰) is similar to the seasonal variation in
average modern δ18Omw values in western Israel (5.2‰), whereas the average amplitude
of δ18Obioapatite values (2.3‰) is about half that.
Local Vegetation and Meteoric Water
The wild vegetation surrounding the three archaeological sites is composed
mainly of trees, shrubs, grasses and herbaceous plants, and has δ13C values ranging from
-32.3‰ to -24.0‰ (see Appendix 3). The δ13C values are indicative of C3 plants, as are
expected in a Mediterranean-type climate. Modern δ18Omw values in western Israel range
from +3.3‰ to -8.5‰ (Fig. 4), with a long-term weighted annual mean of -5.0‰
(IAEA/WMO, 2004). The seasonal variation in average δ18Omw values is ~5.2‰. δDmw
and δ18Omw values from this region fall along a slope of 5.5 (on a δD vs δ18O plot),
indicating that significant evaporation occurs during rainfall (Dansgaard, 1964). The
long-term average annual relative humidity in the region is ~64% (Meterological Office,
1983), and the weighted mean deuterium (d) excess for western Israel is ~18.6
(IAEA/WMO, 2004), which is similar to the calculated Eastern Meteoric Water Line d
excess of ~22 (Gat & Carmi, 1970). This high d excess suggests that humidity is
relatively low in the vapor source region.
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Discussion
Inter-Tooth Sampling: Evidence of Niche Separation and Increasing Aridity
Both Dama and Gazella are characterized as mixed feeders, consuming either
browse or graze depending on seasonal availability (Chapman & Chapman, 1975; Martin,
2000), and we assume here that past feeding behavior is broadly analogous to modern.
Although δ13C values of modern vegetation in western Israel exhibit a range of more than
8‰ (see Appendix 3), and speleothem δ13C records from the region show a range of 12‰
(Frumkin et al., 2000), average Dama and Gazella δ13Cbioapatite values have distinct ranges
of only ~2‰ (Fig. 3a). The average δ13Cbioapatite values of Dama and Gazella range from -
10 to -12‰ and -12 to -14‰, respectively, and show very little change over time. We
interpret this pattern to be indicative of consistent niche separation between Dama and
Gazella, despite gradually increasing aridity and glacial-interglacial climate fluctuations.
It is also evident that the niche for Gazella was available at the time of the species’
sudden influx into the Levant region ~200 ka BP. Dama and Gazella appear to have been
selectively consuming discrete types of C3 vegetation throughout both glacial and
interglacial periods, possibly suggesting that climate change did not drastically alter the
types of vegetation present in the region.
In contrast with the δ13Cbioapatite values, the δ18Obioapatite record is likely climate-
dependent (Fig. 3b). Average Dama and Gazella δ18Obioapatite values increase by ~4‰
over the past 350 ka, indicating a trend toward gradually increasing aridity over time. The
suggestion of increasing aridity in this region is supported by isotopic (Shahack-Gross et
al., 1999), pollen (Albert et al., 2003) and microfauna (Tchernov, 1994) evidence. In
addition, because Gazella typically pant (as opposed to sweating) and select plant leaves
15
over stems, their δ18Obioapatite values tend to be slightly elevated compared to Dama
(Kohn, 1996).
It appears that vegetation communities in the Levant were relatively stable over
the past 350 kyr, and that aridity increased only gradually over time. Hence, it is likely
that the sudden dispersal of Gazella from the Afro-Arabian biotic province into the
Levant region ~200 ka BP is independent of any sort of drastic climate or environmental
change. It is possible that the increasing proportion of Gazella in the archaeological
record after ~200 ka BP is due mainly to changes in human subsistence behavior and prey
selection, and less to climate-driven environmental changes.
The most recent glaciation (Oxygen Isotope Stage 2) is well represented by the
ungulate tooth time series. Average δ18Obioapatite values increase ~30 ka BP, and
subsequently decrease by 2‰ by the end of the glacial interval ~15 ka BP. These data
correspond strongly with speleothem δ18O (δ18Ocalcite) records from the region (Bar-
Matthews et al., 1999; Frumkin et al., 1999) (Fig. 5), which indicate that δ18Ocalcite values
are higher during glacial periods and lower during interglacial periods. Frumkin et al.
(1999) explain this unusual isotopic pattern by postulating that excess evaporation and
limited circulation in the eastern Mediterranean Sea during glacial periods causes higher
δ18O values of sea surface water, and hence, higher δ18Omw and δ18Ocalcite values across
the adjacent Levant.
16
Inter-Tooth Sampling: δ18Omw Reconstruction and δ18Obioapatite Modeling
To estimate past variation of mean δ18Omw values in western Israel, we extrapolate
from the Frumkin et al. (1999) Jerusalem Cave δ18Ocalcite record using the following
expression from Kim and O’Neil (1997):
(2)
where α is the fractionation factor, T is the temperature in Kelvin, and
. (3)
A rough approximation of past mean δ18Omw values can be reconstructed using measured
glacial and interglacial δ18Ocalcite values (Frumkin et al., 1999) and the modern mean
annual temperature (MAT) of 17 °C (IAEA/WMO, 2004) (Fig. 5). In addition, we use an
estimated glacial-interglacial ∆MAT of about 5‰, and we take into account a factor of
4‰ to account for limited circulation and excess evaporation of the Mediterranean Sea
during glacial periods (Frumkin et al., 1999). If δ18O values of herbivore drinking water
(δ18Odw) are assumed to be comparable to average δ18Omw values, this basic
reconstruction can be used to compare the enrichment factors between estimated δ18O
values of ingested waters and both measured and modeled δ18Obioapatite values.
Several predictive models of mammalian oxygen isotope composition have been
developed (Luz & Kolodny, 1985; Ayliffe & Chivas, 1990; Luz et al., 1990; Bryant &
Froelich, 1995; Kohn, 1996), and in Figure 5 we compare our average observed
δ18Obioapatite values to the outputs from these models. The best fit to our data is attained
with the Bryant & Froelich (1995) model, which uses body mass-dependent scaling
equations. Although the scaling methods used in this model are not sensitive enough to
17
predict the measured species-specific differences in Dama and Gazella δ18Obioapatite
values, the model predicts average herbivore δ18Obioapatite values very close to what we
observe.
It is surprising that the Kohn (1996) model, which is based on a genus-specific
approach, underpredicts many of the Gazella δ18Obioapatite values by up to 6‰.
Additionally, the Luz & Kolodny (1985) model, which is based on small herbivores,
underpredicts the observed δ18Obioapatite values up to 8‰. Both the Kohn (1996) and the
Bryant & Froelich (1995) models use similar sources of data and take into account the
oxygen isotope fractionations in consumed vegetation, but the Bryant & Froelich (1995)
study relies on representative proportions of oxygen fluxes and fractionations for all
species, whereas the Kohn (1996) study incorporates genus-specific differences in animal
diet, respiratory water vapor gain and loss, and heat regulation and waste-loss
mechanisms. Kohn (1996) acknowledges that the model underpredicts δ18Obioapatite values
for antelope genera such as Gazella, and that these predictions become consistent with
measured values once additional transcutaneous water vapor loss is included in the model
parameters. However, in contrast with the Bryant & Froelich (1995) and Luz & Kolodny
(1985) models, Kohn’s genus-specific model does not predict the full extent of glacial-
interglacial δ18Obioapatite variation.
Our data suggest that average Pleistocene Dama and Gazella tooth enamel is enriched
(εbioapatite-water) on the order of 29-37‰ with respect to reconstructed δ18Omw values, a
much larger enrichment factor than that predicted by recent physiologic models (~26‰,
Bryant et al., 1996b). Gazella exhibits a slightly greater εbioapatite-water than Dama, which is
especially evident during the last glacial period (Fig. 5). These results point toward
18
significant species-specific enrichment of δ18Obioapatite values, in which differing
physiologies (panting vs. sweating, total water turnover, proportions of oxygen fluxes)
and diets (a preference for leaves or stems) play a large role. This conclusion is in strong
agreement with the recent work of Levin et al. (2006).
Intra-tooth Sampling: Evidence of Seasonal Variation
Large intra-tooth variation in both δ13Cbioapatite and δ18Obioapatite values is apparent
in our dataset (see Appendices 1 & 2), which we interpret as reflecting seasonal changes
in climate and diet composition during the time of tooth formation. Although Dama and
Gazella bioapatite does not likely capture the full amplitude of annual variation of
δ13Cvegetation and δ18Omw values, it does provide a minimum estimate of past seasonality.
There are at least four potential sources of isotopic attenuation in tooth enamel:
residence time of carbon and oxygen in an animal, environmental averaging of isotopic
extremes, resolution of the microsampling procedure, and the duration of tooth
mineralization. Residence times of carbon and oxygen in a large herbivore are on the
order of several weeks (Kohn et al., 2002; Ayliffe et al., 2004), implying that isotopic
extremes in the environment will be somewhat smoothed in the bioapatite record. Some
attenuation of the isotopic values will also occur because of environmental averaging,
and the microsampling procedure. This attenuation is easily observed when comparing
the average intra-tooth δ18Obioapatite variation of 2.3‰ to the modern δ18Omw amplitude of
5.2‰ (Fig. 4). Even the maximum intra-tooth range of δ18Obioapatite values (4.1‰) is
somewhat less than the observed seasonal variation in average modern δ18Omw values in
19
western Israel. Lastly, intra-tooth isotopic variation may also appear attenuated if the
third molar completes mineralization over a period of less than ~6 months.
Unfortunately, the length of time represented in Dama and Gazella teeth is
difficult to constrain. Kohn et al. (1998) have estimated that 1.9 mm of enamel is
mineralized per week in African gazelle third molars. Fricke and O’Neil (1996) find
slightly lower rates of enamel mineralization in other ungulates, from 0.4-0.8 mm/week.
Average Dama and Gazella M3s in this study are ~15 mm in length, implying that
complete enamel mineralization takes somewhere on the order of 8 to 38 weeks. Hence,
less than an annual cycle is likely represented in a single tooth, and the intra-tooth
measurements can be interpreted as a minimum estimate of past seasonality (Fig. 6).
The consumption of seasonally available types of vegetation may account for
intra-tooth variation of δ13Cbioapatite values. Additionally, most semiarid plant species
naturally undergo a 1-3‰ change in carbon isotope values during the growing season
(Ehleringer et al., 1992). Taking into account the discrete niches for Dama and Gazella
discussed above, the measured intra-tooth δ13Cbioapatite ranges of 0.4-3.6‰ are likely due
to consumption of a select variety of seasonally available plants. On average, Gazella
show slightly more variation in intra-tooth δ13Cbioapatite values than do Dama (1.2‰ as
opposed to 0.9‰). This may be due either to the fact that most Gazella teeth are higher-
crowned and thus capture more isotopic variability, or to the possibility that Gazella were
consuming a somewhat more varied diet. Overall, there are few noticeable patterns in the
intra-tooth δ13Cbioapatite data, and the range of δ13Cbioapatite variance stays remarkably
similar over time. It is interesting to note, however, that most of the Aurignacian-age
Dama and Gazella teeth from Hayonim Cave display a high degree of covariance
20
between the δ13Cbioapatite and δ18Obioapatite values that teeth from other time periods do not
(see Appendix 2).
Intra-tooth variation in δ18Obioapatite values ranges from 0.3-4.1‰, and is likely due
to seasonal changes in rainfall and humidity. Leaf water can be evaporatively enriched in
18
O by 10-25‰ in comparison to stem and local meteoric water (Dongmann et al., 1974),
and may significantly influence δ18Obioapatite values. If conditions become more arid or if
herbivores are ingesting a greater proportion of leaves, the intra-tooth δ18Obioapatite range is
likely to increase. It has been observed that modern gazelle populations in western Israel
are independent of standing water, and tend to meet their water requirements by ingesting
vegetation that is coated in morning dew, by selecting plants that have high water
content, and by feeding during cooler times of the day (Martin, 2000). It is likely that
deer in this region follow similar practices (Chapman & Chapman, 1975), and that both
species behaved in a comparable manner in the past.
The average variation in intra-tooth δ18Obioapatite values for Dama and Gazella for
most time periods is 2-2.4‰, but Natufian gazelles show average δ18Obioapatite variation of
only 1.3‰. This pattern may be explained by a proposed cooler and/or wetter climate
during the Natufian period in western Israel (Shahack-Gross et al., 1999), which would
imply that it was less arid and that Gazella were consuming vegetation with lower δ18Olw
values. However, given that the overall trend in δ18Obioapatite values seems to be one of
gradually increasing aridity, it is unclear exactly why the Natufian intra-tooth values are
so much lower than average.
21
Intra-tooth Sampling: Birthing Seasons
Previous studies have suggested that substantial intra-tooth isotope variation in an
herbivore population may be explained by multiple birthing peaks and subsequent
bioapatite growth and mineralization throughout the year (Shahack-Gross et al., 1999;
Balasse et al., 2003). Because climatic and environmental factors seasonally influence
δ18Omw values, the patterns and offset of intra-tooth δ18Obioapatite curves among a species
can be examined in order to investigate different birthing seasons. The annual birthing
season window is generally most restricted and consistent for mammals living at high
latitudes, but it expands at lower latitudes in a given species. Although exceptions exist,
we can expect ungulate birthing seasons to be variable in the study area because of its
position at ~32-33° N latitude. Of interest over long time spans are possible changes in
birthing synchrony among species that might reflect responses to changes in the
seasonality of the environment. Time-averaging is a potential obstacle to such an
analysis, but generalized differences among periods may still be informative.
Gazella in western Israel have been observed to reproduce year-round with semi-
annual birthing peaks during April-June and November (Martin, 2000), whereas Dama
typically have only one birthing peak per year in May-June (Chapman & Chapman,
1975). The patterns of intra-tooth variation in our δ18Obioapatite data are consistent with
these modern observations (Fig. 7a and 7b). Gazelles appear to have multiple birthing
seasons, as shown by two fairly distinct groupings of intra-tooth isotope curves. Fallow
deer likely have only one birthing season, as most intra-tooth isotope curves have a
similar shape and are offset very little.
22
On average, Dama exhibit slightly more variability (2.4‰) in intra-tooth
δ18Obioapatite values than do Gazella (2.0‰). Dama M3 eruption times are between 20-26
weeks of age (Chapman & Chapman, 1975), and Gazella M3 eruption times are from 40-
50 weeks of age (Davis, 1980). Accordingly, M3s in Dama and in gazelles born during
the November birthing peak would undergo mineralization during the cool season when
more moisture is available and δ18Olw values are lower. Gazelles born in the early
summer would have M3s mineralizing during the warmer season when leaf water is more
enriched in 18O. There is typically a greater propensity for larger ranges of δ18Olw values
during the cool season (shown by the Dama teeth), as compared to during the warm
season when most δ18Olw values are very high and will likely show little variation.
However, the differences in intra-tooth δ18Obioapatite variability between the two species
may also be due specific diet preferences and feeding habits.
Concluding Remarks
Our study showed for the first time that pretreatment procedures for enamel
microsamples of various ages are largely unnecessary. Although neither the mean
δ18Obioapatite nor δ13Cbioapatite values of the unpretreated and fully pretreated fossil tooth
enamel were statistically within error at the 95% confidence level, the differences
between the two sets of mean δ18Obioapatite and δ13Cbioapatite values were relatively small
when compared to other sources of uncertainty in the study. Hence, a careful
microsampling methodology may preclude any need to pretreat enamel samples.
Through stable isotope analysis of tooth enamel it is possible to gain insight on
how and to what degree mammals responded to broad-scale changes in climate and
23
environment over time, as well as to obtain glimpses into past seasonal variability. These
data may also help test hypotheses about whether changes in climate can explain
observed patterns in the faunal records. Results from both inter- and intra-tooth analyses
can be used as a framework against which to better understand the Paleolithic
archaeological and human evolutionary record in the Levant. Although intra-tooth
isotopic variation in δ18Obioapatite is attenuated with respect to probable seasonal variation
in δ18Omw, intra-tooth measurements permit a minimum estimation of that seasonal
variability, and can be used to reconstruct such patterns as birthing seasonality. In
paleoclimate studies such as this, it is essential to analyze enamel from more than one
species, as herbivores have specific diet preferences and physiologies that uniquely
influence the isotope signature recorded in their bioapatite.
It does not appear that climate change was the driving factor behind the rather
sudden dispersal of Gazella from the Afro-Arabian biotic province into the
Mediterranean hills of the Levant region ~200 ka BP. Rather than an abrupt change in
climate and environmental conditions around 200 ka BP, our data instead suggest a trend
of gradually increasing aridity over time. Additionally, both Gazella and Dama occupied
distinct, unchanging niches for the entirety of the record examined by this study,
indicating that perhaps vegetation communities in the region varied relatively little
between glacial and interglacial periods.
Comparisons of measured δ18Obioapatite values with reconstructed δ18Omw values
suggest that average Dama and Gazella tooth enamel is enriched (εbioapatite-water) on the
order of 29-37‰, a much larger enrichment factor than that predicted by recent
physiologic models (26‰, Bryant et al., 1996b). These findings point toward significant
24
species-specific enrichment of δ18Obioapatite values, which is likely due to particular diet
preferences and physiologic factors (Levin et al., 2006). The utility of the 26‰
enrichment factor should be reevaluated in the future and caution should be exercised
when relying on it in further research.
Acknowledgements
We thank the Israel Antiquities Authority for permission to export ungulate tooth
samples for destructive analysis, and we are grateful to Ofer Bar-Yosef, Rivka
Rabinovich, Guy Bar-Oz, Natalie Munro and Steven Kuhn for permission, assistance or
both in obtaining the samples from the faunal collections of Qesem Cave, Hayonim Cave
and Meged Rockshelter. We also thank Nuno Bicho for permission to sample the tooth
from Val Boi, Portugal for the pretreatment experiment. This research was funded by a
National Science Foundation Grant (BCS-0410654) to M.C.S. and a fellowship from the
National Science Foundation’s IGERT Program in Archaeological Sciences (DGE-
0221594) to J.C.R. Microsampling advice from Naomi Levin was fundamental to this
study, and is very much appreciated. Conversations with Stanley Ambrose and Matthew
Sponheimer also contributed greatly to the improvement of this project.
25
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Zazzo, A., Balasse, M.& Patterson, W.P. (2005). High-resolution δ13C intratooth profiles in bovine enamel:
Implications for mineralization pattern and isotopic attenuation. Geochimica et Cosmochimica Acta 69,
3631-3642.
Zohary, M. (1973). Geobotanical Foundations of the Middle East. Stuttgart: Springer Verlag.
Zohary, M. & Feinbrun-Dothan, N. (1966-1986). Flora Palaestina, Volumes 1-4. Jerusalem: Publications of
the Israel Academy of Sciences and Humanities.
30
Table 1
Age estimates for Hayonim, Meged and Qesem
* * *
Period Site, Culture & Layer Age Range (ka BP)
Epipaleolithic Hayonim Cave, Natufian (layer B) a,b
11-13
Hayonim Cave, Kebaran (layer C) b
14-17
Meged Rockshelter, Kebaran (<200 cm) c
18-19
Upper Paleolithic Hayonim Cave, Aurignacian (layer D) a,d
26-28
Middle Paleolithic Hayonim Cave, Mousterian Units 1-2 (layer E) e
70-100
Hayonim Cave, Mousterian Unit 3 (layer E) e,f,g
~150
Hayonim Cave, Mousterian Unit 4 (layer E) e,g
~170
Hayonim Cave, Mousterian Unit 5-6 (layer E) e,f,g
~200
Hayonim Cave, Mousterian Unit 7 (layer F) e,g
200-215
Lower Paleolithic Qesem Cave, Achuelo-Yabrudian (layer 2) h
200-230
Qesem Cave, Achuelo-Yabrudian (layer 4) h
300-350
Age estimates are from aBar-Yosef, 1991; bHousley, 1994; cKuhn et al., 2004; dPhillips, 1994; eSchwarcz
and Rink, 1998; fValladas et al., 1998; gRink et al., 2004; hBarkai et al., 2003. *See text for specific dating
methods used and further explanation of accepted cave chronologies. Note: Epipaleolithic and Upper
Paleolithic radiocarbon dates are uncalibrated.
31
Table 2
Pretreatment experiment data
Modern tooth - Equus kiang (Tibet) Fossil tooth - Equus caballus ~28 ka (Val Boi, Portugal)
type of δ13C PDB C STD δ18O PDB O STD type of δ13C PDB C STD δ18O PDB O STD
pretreatment sample ID measured (‰) (σ) measured (‰) (σ) pretreatment sample ID measured (‰) (σ) measured (‰) (σ)
no pretreatment 030605-9A1 -12.04 0.034 -8.17 0.075 no pretreatment ISO820A1 -11.75 0.013 -0.38 0.075
"a" 030605-9A2 -12.05 0.019 -8.17 0.088 "a" ISO820A2 -11.67 0.040 -0.59 0.050
030605-9A3 -12.11 0.037 -8.32 0.042 ISO820A3 -11.70 0.023 -0.40 0.051
030605-9A4 -12.04 0.035 -8.17 0.082 ISO820A4 -11.66 0.057 -0.48 0.061
030605-9A5 -12.04 0.012 -8.24 0.037 ISO820A5 -11.70 0.036 -0.42 0.037
030605-9A6 -12.02 0.028 -8.27 0.045 ISO820A6 -11.65 0.034 -0.47 0.122
030605-9A7 -11.99 0.061 -8.21 0.067 ISO820A7 -11.59 0.024 -0.56 0.055
030605-9A8 -12.03 0.029 -8.20 0.082 ISO820A8 -11.63 0.067 -0.57 0.045
030605-9A9 -12.02 0.020 -8.24 0.064 ISO820A9 -11.59 0.043 -0.51 0.101
030605-9A10 -12.01 0.030 -8.22 0.056 ISO820A10 -11.65 0.022 -0.63 0.020
wt. mean: -12.04 -8.24 wt. mean: -11.69 -0.55
error wt. mean: 0.0073 0.0177 error wt. mean: 0.008 0.0136
95% confidence: -12.04±0.005 -8.24±0.01 95% confidence: -11.69±0.01 -0.55±0.01
bleach only 030605-9B1 -12.13 0.021 -7.65 0.050 bleach only ISO820B1 -11.76 0.032 0.36 0.091
"b" 030605-9B2 -12.17 0.026 -7.39 0.058 "b" ISO820B2 -11.76 0.016 0.07 0.028
030605-9B3 -12.17 0.021 -7.70 0.070 ISO820B3 -11.75 0.052 0.46 0.097
030605-9B4 -12.20 0.022 -7.49 0.012 ISO820B4 -11.72 0.028 0.16 0.079
030605-9B5 -12.20 0.020 -7.75 0.110 ISO820B5 -11.81 0.020 -0.03 0.039
wt. mean: -12.17 -7.50 wt. mean: -11.77 0.08
95% confidence: -12.17±0.01 -7.50±0.01 95% confidence: -11.77±0.01 0.08±0.03
bleach + acetic 030605-9C1 -12.22 0.040 -8.24 0.098 bleach + acetic ISO820C1 -11.62 0.025 -0.24 0.100
"c" 030605 9C2 -12.24 0.026 -8.21 0.070 "c" ISO820C2 -11.59 0.039 -0.13 0.041
020605 9C3 -12.19 0.043 -8.26 0.088 ISO820C3 -11.68 0.030 -0.22 0.048
030605 9C4 -12.23 0.038 -8.26 0.071 ISO820C4 -11.63 0.021 -0.19 0.047
030605 9C5 -12.20 0.020 -8.21 0.038 ISO820C5 -11.63 0.030 -0.13 0.077
wt. mean: -12.22 -8.23 wt. mean: -11.63 -0.18
acetic only 030605-9D1 -12.19 0.020 -7.50 0.123 acetic only ISO820D1 -11.70 0.046 0.95 0.050
"d" 030605 9D2 -12.22 0.037 -7.59 0.093 "d" ISO820D2 -11.75 0.013 0.64 0.061
020605 9D3 -12.22 0.033 -7.61 0.051 ISO820D3 -11.73 0.045 0.86 0.034
030605 9D4 -12.25 0.036 -7.41 0.072 ISO820D4 -11.71 0.015 0.66 0.089
030605 9D5 -12.20 0.014 -7.48 0.098 ISO820D5 -11.73 0.021 0.86 0.037
wt. mean: -12.20 -7.54 wt. mean: -11.73 0.84
95% confidence: -12.20±0.01 -7.54±0.04 95% confidence: -11.73±0.01 0.84±0.03
32
Table 2 Continued
Pretreatment experiment data
Fossil tooth - Mammuthus columbi 11-12 ka (Las Vegas, NV area) Fossil tooth - Gomphothere ~11 Ma (Pascalar, Turkey)
type of δ13C PDB C STD δ18O PDB O STD type of δ13C PDB C STD δ18O PDB O STD
pretreatment sample ID measured (‰) (σ) measured (‰) (σ) pretreatment sample ID measured (‰) (σ) measured (‰) (σ)
no pretreatment MAM A1 -7.21 0.024 -9.73 0.091 no pretreatment GOM A1 -11.04 0.031 -6.14 0.040
"a" MAM A2 -7.21 0.038 -9.64 0.075 "a" GOM A2 -10.98 0.029 -6.36 0.054
MAM A3 -7.18 0.025 -9.75 0.033 GOM A3 -11.05 0.030 -6.01 0.024
MAM A4 -7.14 0.047 -9.73 0.066 GOM A4 -11.00 0.047 -6.37 0.052
MAM A5 -7.15 0.027 -9.80 0.051 GOM A5 -11.02 0.017 -6.13 0.060
wt. mean: -7.18 -9.75 wt. mean: -11.02 -6.12
95% confidence: -7.18±.002 -9.75±0.03 95% confidence: -11.02±0.01 -6.12±0.02
bleach only MAM B1 -7.83 0.033 -8.75 0.064 bleach only GOM B1 -11.10 0.029 -5.27 0.120
"b" MAM B2 -7.77 0.020 -8.55 0.035 "b" GOM B2 -11.11 0.026 -5.56 0.053
MAM B3 -7.75 0.027 -8.86 0.081 GOM B3 -11.09 0.019 -5.45 0.070
MAM B4 -7.77 0.049 -8.53 0.042 GOM B4 -11.06 0.026 -5.43 0.021
MAM B5 -7.73 0.017 -8.69 0.078 GOM B5 -11.10 0.038 -5.49 0.049
wt. mean: -7.76 -8.61 wt. mean: -11.09 -5.45
bleach + acetic MAM C1 -7.68 0.044 -9.74 0.052 bleach + acetic GOM C1 -11.24 0.053 -5.82 0.040
"c" MAM C2 -7.69 0.042 -9.48 0.068 "c" GOM C2 -11.20 0.039 -5.90 0.073
MAM C3 -7.67 0.028 -9.65 0.042 GOM C3 -11.19 0.011 -5.80 0.102
MAM C4 -7.73 0.011 -9.48 0.048 GOM C4 -11.21 0.033 -5.81 0.059
MAM C5 -7.64 0.025 -9.67 0.060 GOM C5 -11.21 0.043 -5.77 0.048
wt. mean: -7.71 -9.61 wt. mean: -11.20 -5.81
acetic only MAM D1 -7.75 0.008 -9.90 0.038 acetic only GOM D1 -11.23 0.009 -6.00 0.050
"d" MAM D2 -7.76 0.022 -9.81 0.076 "d" GOM D2 -11.23 0.035 -6.00 0.048
MAM D3 -7.77 0.048 -9.87 0.091 GOM D3 -11.23 0.020 -5.75 0.079
MAM D4 -7.76 0.034 -9.70 0.055 GOM D4 -11.26 0.030 -5.67 0.087
MAM D5 -7.71 0.031 -9.74 0.045 GOM D5 -11.27 0.010 -5.45 0.082
wt. mean: -7.75 -9.81 wt. mean: -11.24 -5.87
33
Figure Captions
Figure 1. Map showing locations of Qesem Cave, Hayonim Cave and Meged Rockshelter
in the Levant region of Israel. Jerusalem Cave, the source of the Frumkin et al. (1999)
speleothem study, is also shown.
Figure 2. Comparison of (a) average δ13Cno pretreatment versus δ13Cfull pretreatment and (b)
average δ18Ono pretreatment versus δ18Ofull pretreatment values. Straight lines are 1:1 lines. Error
bars for each point are smaller than the size of the symbol.
Figure 3. Inter-tooth comparison of (a) average δ13Cbioapatite values and (b) average
δ18Obioapatite values. Trendlines for both Dama and Gazella are also shown. Error bars
represent the range of measured intra-tooth isotopic variability, and shaded boxes indicate
cooler, glacial periods during the 350-kyr record. Data are described in detail in the text.
Figure 4. Modern climate data from the Bet Dagan weather station near Tel Aviv, Israel.
Graphs show (a) mean monthly precipitation from 1960-2001, (b) mean monthly
temperature from 1960-1979, (c) mean monthly δ18Omw measurements from 1960-2001,
and (d) δDmw versus δ18Omw measurements from 1960-2001 compared to the global
meteoric water line (GMWL). Modern climate data are used as an analog for probable
average interglacial conditions in the Levant region.
Figure 5. (a) Average δ18Obioapatite values compared against the predictive model curves of
Bryant & Froelich (1995), Kohn (1996) and Luz & Kolodny (1985). The Bryant &
Froelich (1995) predictive model provides the best fit with our data. Predictive models
were calculated using (b) reconstructed δ18Omw values as estimated from speleothem δ18O
measurements from Jerusalem Cave (Frumkin et al., 1999). (c) The SPECMAP stacked
δ18O curve (after Imbrie et al., 1984) defining glacial (shaded) and interglacial periods.
Enlarged box in upper left corner of figure shows a portion of the speleothem δ18O curve
from Frumkin et al. (1999) and measured δ18Obioapatite values from the last glacial period,
confirming that δ18O values increase during glacial periods (likely due to decreased
Mediterranean circulation and increased evaporation; see text for further explanation).
Figure 6. Sequential sampling of enamel along the tooth provides a record of intra-tooth
δ18Obioapatite and δ13Cbioapatite variation. The neck of the tooth is defined as the point where
the roots meet the enamel. Shown is sample ISO-667 (Gazella UM3 from Hayonim
Cave, ~170 ka BP). Almost a full seasonal cycle is represented in this tooth.
Figure 7. Birthing seasonality of (a) Dama (from Qesem Cave, 350-300 ka BP) and (b)
Gazella (from Hayonim Cave, 200-150 ka BP) can be interpreted from intra-tooth
variation in δ18Obioapatite values. Dama appear to have one birthing season (intra-tooth
curves are similarly shaped), whereas Gazella likely have two (distinguished by two
distinct groupings of intra-tooth curves with heavy and light connecting lines).
34
35°E
33°N
Haifa
Hayonim,
N Meged
Tel Aviv
32°N
Qesem
Jerusalem
31°N
ISRAEL
EGYPT JORDAN
30°N
50 km
Figure 1
35
-6 32
a b
30
δ13C no pretreatment
δ18O no pretreatment
-8
28
-10 26
Modern kiang Modern kiang

24
Equus caballus Equus caballus
-12 Mammuthus Mammuthus
Gomphothere 22 Gomphothere
36
1:1 line 1:1 line

-14 20
-14 -12 -10 -8 -6 20 22 24 26 28 30 32
δ13C full pretreatment δ18O full pretreatment
Figure 2
-5 37
-6 a y = 0.0004x - 11.116 36 y = -0.0024x + 31.305 b
2
-7 R2 = 0.015 35
R = 0.0167
-8 34
δ O bioapatite(‰)
δ C bioapatite (‰)
-9 33
-10 32
-11 31
-12 30
-13 29
18
13
-14 28
37
-15 27
Dama Dama
-16 Gazella 26 Gazella
y = -7E-05x - 12.847 y = -0.0047x + 30.435
-17 2 Capra 25 Capra
R = 7E-05 Bos Bos R2 = 0.371
-18 24
0 50 100 150 200 250 300 350 400 0 50 100 150 200 250 300 350 400
time (ka BP) time (ka BP)
Figure 3
160 0
140
a c
-1
precipitation (mm)
120
-2
δ18Omw (‰)
100
80 -3
60
-4
40
-5
20
0 -6
J F M A M J J A S O N D J F M A M J J A S O N D
month month
38
30 30
b 20 d
25
10
temperature (C)
20 0
δDmw (‰)
y = 5.49x + 6.22
-10
15 -20
-30
10
-40
-50
5 WL
-60 GM
0 -70
J F M A M J J A S O N D -10 -8 -6 -4 -2 0 2 4
month δ18Omw (‰)
Figure 4
34
33 a. measured and modeled enamel δ18O values
32
34
(‰)
31
bioapatite
30
32
δ18O bioapatite (‰)

29
28 Bryant & Froelich, 1995 (gazelle)
27 30
δ O
18
26 Bryant & Froelich, 1995 (deer)

25
24 28
0 20 40 Kohn, 1996 (gazelle)
time (ka BP)
26
Luz and Kolodny, 1985 (rats)
24
b. reconstructed δ18O meteoric water from speleothems
2
Soreq Cave, Bar-Matthews et al, 1999
39
δ18O mw (‰)
-2
-4
Jerusalem Cave, Frumkin et al, 1999
-6
2.5 c. SPECMAP -8
stacked δ18O (‰)
1.5
0.5
-0.5
-1.5
-2.5
0 100 200 300 400
time (ka BP)
Figure 5
base
crown
time
31.5
summer
31.0
30.5
30.0
29.5
29.0 winter
28.5
-13.1
summer -13.2
δ13C bioapatite (‰)

-13.3
-13.4
-13.5
-13.6
winter -13.7
-13.8
22 20 18 16 14 12 10 8 6 4 2 0
distance from neck (mm)
Figure 6
40
33 34
32 33
31

32
30
31
29
30
28
29
27
41
26 28
25 27
13 12 11 10 9 8 7 6 5 4 3 2 1 0 22 20 18 16 14 12 10 8 6 4 2 0
distance from neck (mm) distance from neck (mm)
Figure 7
Appendix 1
Intratooth Microsampling Data
δ13C PDB δ18O PDB Distance
Tooth Estimated (measured, C (measured, δ18O O from neck
Sample ID Species element age range ‰) STD ‰) SMOW (‰) STD (mm)
QESEM
QC 411A Dama LM3 300-350ka -10.42 0.039 -1.01 29.87 0.072 12.0
QC 411B -10.56 0.036 -1.03 29.84 0.112 10.3
QC 411C -10.81 0.047 -1.23 29.64 0.041 9.0
QC 411D -10.84 0.026 -2.11 28.73 0.096 7.8
QC 411E -10.81 0.020 -2.61 28.22 0.080 6.6
QC 411F -11.14 0.034 -2.80 28.02 0.072 5.3
QC 411G -11.20 0.037 -2.75 28.07 0.075 4.1
QC 411H -11.40 0.044 -2.82 28.00 0.054 2.9
QC 411I -11.59 0.034 -1.91 28.94 0.075 1.8
QC 411J -11.85 0.027 -2.40 28.43 0.109 0.7
average: -11.06 28.78
QC 417A Dama LM3 300-350ka -10.85 0.009 -1.81 29.04 0.022 11.2
QC 417B -10.59 0.028 -1.82 29.03 0.040 10.0
QC 417C -10.37 0.047 -2.49 28.34 0.057 8.8
QC 417D -10.33 0.010 -3.08 27.73 0.041 7.8
QC 417E -10.25 0.009 -2.66 28.16 0.039 6.5
QC 417F -10.11 0.029 -2.38 28.45 0.082 5.7
QC 417G -10.11 0.026 -2.28 28.56 0.114 4.5
QC 417H -9.99 0.038 -1.66 29.20 0.052 3.2
QC 417I -10.42 0.041 -1.20 29.67 0.101 2.3
QC 417J -10.91 0.017 -1.02 29.86 0.035 1.1
average: -10.39 28.80
QC 425A Dama LM3 300-350ka -11.35 0.057 -3.06 27.75 0.063 7.3
QC 425B -11.40 0.022 -3.14 27.67 0.048 6.1
QC 425C -11.52 0.023 -2.91 27.91 0.029 5.2
QC 425D -11.13 0.046 -3.01 27.80 0.029 4.0
QC 425E -11.28 0.018 -2.81 28.01 0.065 2.9
QC 425F -11.43 0.016 -2.51 28.32 0.068 1.8
QC 425G -11.08 0.028 -2.20 28.64 0.028 0.7
average: -11.31 28.01
QC 430A Dama LM3 300-350ka -11.10 0.038 1.11 32.05 0.014 9.8
QC 430B -11.36 0.051 1.57 32.53 0.074 8.5
QC 430C -10.93 0.012 -0.31 30.59 0.044 7.4
QC 430D -11.37 0.020 0.33 31.25 0.070 6.6
QC 430E -11.43 0.018 0.46 31.39 0.038 5.2
QC 430F -11.00 0.040 -0.54 30.35 0.031 4.3
QC 430G -10.73 0.018 -1.92 28.93 0.057 3.5
QC 430H -11.58 0.024 -1.65 29.20 0.047 2.4
QC 430I -11.10 0.017 -2.31 28.53 0.027 1.2
average: -11.18 30.54
42
QC426A Dama LM3 300-350ka -10.72 0.023 -1.00 29.87 0.046 10.7
QC426B -10.85 0.019 -1.79 29.06 0.025 9.3
QC426C -10.65 0.020 -2.45 28.38 0.035 7.9
QC426D -10.67 0.022 -2.67 28.16 0.093 6.4
QC426E -11.09 0.045 -3.20 27.61 0.040 5.0
QC426F -11.22 0.026 -3.32 27.48 0.057 3.6
QC426G -11.45 0.021 -2.72 28.10 0.035 2.5
QC 426H -11.77 0.032 -2.28 28.55 0.034 1.1
average: -11.05 28.40
QC 432A Dama LM3 300-350ka -11.00 0.039 -0.81 30.07 0.045 8.7
QC 432B -11.02 0.033 -1.22 29.65 0.035 7.4
QC 432C -10.79 0.039 -2.05 28.80 0.030 6.5
QC 432D -11.03 0.032 -2.47 28.36 0.015 5.6
QC 432E -10.95 0.021 -2.93 27.88 0.058 4.8
QC 432F -11.25 0.018 -3.02 27.79 0.057 3.5
QC 432G -11.20 0.021 -3.23 27.58 0.037 2.4
QC 432H -10.77 0.033 -2.30 28.53 0.018 1.5
average: -11.00 28.58
QC 428A Dama LM3 300-350ka -10.78 0.010 -1.66 29.19 0.048 8.2
QC 428B -10.52 0.043 -2.17 28.66 0.037 7.3
QC 428C -10.55 0.018 -2.53 28.30 0.099 6.2
QC 428D -10.55 0.018 -3.44 27.36 0.081 5.0
QC 428E -10.82 0.031 -4.11 26.67 0.029 4.1
QC 428F -10.98 0.021 -4.31 26.47 0.053 3.3
QC 428G -10.67 0.036 -4.58 26.18 0.093 2.3
QC 428H -10.50 0.046 -3.45 27.35 0.125 1.0
average: -10.67 27.52
QC 438A Dama LM3 200-230ka -11.57 0.005 0.08 30.99 0.035 10.8
QC 438B -11.41 0.023 0.67 31.60 0.081 9.2
QC 438C -11.44 0.015 0.41 31.33 0.115 8.0
QC 438D -11.54 0.030 -0.14 30.76 0.045 6.6
QC 438E -11.78 0.042 -0.90 29.98 0.061 5.4
QC 438F -12.20 0.028 -2.02 28.82 0.068 4.6
QC 438G -12.15 0.020 -2.08 28.76 0.084 3.1
QC 438H -12.13 0.037 -1.98 28.87 0.042 1.7
average: -11.78 30.14
QC 442A Dama LM3 200-230ka -10.33 0.029 -0.25 30.65 0.061 11.4
QC 442B -10.41 0.028 -1.13 29.74 0.033 10.1
QC 442C -10.20 0.038 -1.64 29.22 0.045 9.0
QC 442D -10.16 0.035 -1.90 28.95 0.042 7.8
QC 442E -10.28 0.029 -1.91 28.94 0.050 6.9
QC 442F -10.59 0.021 -1.84 29.01 0.036 5.7
QC 442G -10.50 0.033 -1.98 28.87 0.066 4.3
QC 442H -10.73 0.011 -2.06 28.78 0.037 3.5
43
QC 442I -10.73 0.007 -1.76 29.09 0.084 2.3
QC 442J -10.57 0.038 -2.02 28.82 0.037 1.1
average: -10.45 29.21
HAYONIM-E. MIDDLE PALEOLITHIC

ISO 681A Dama UP3 >200,000 -11.12 0.019 -0.20 30.70 0.037 12.9
ISO 681B -10.84 0.035 -0.05 30.86 0.027 11.7
ISO 681C -10.49 0.011 0.00 30.90 0.039 10.6
ISO 681D -10.75 0.016 0.33 31.25 0.080 9.4
ISO 681E -10.25 0.023 -0.14 30.76 0.048 8.1
ISO 681F -10.48 0.022 -0.71 30.18 0.060 7.0
ISO 681G -10.49 0.021 -0.63 30.26 0.033 5.8
ISO 681H -10.56 0.043 -1.10 29.78 0.055 4.9
ISO 681I -10.80 0.027 -1.62 29.23 0.069 3.8
ISO 681J -10.66 0.042 -2.09 28.75 0.067 2.6
ISO 681K -10.61 0.017 -2.43 28.40 0.079 1.2
average: -10.64 30.10
ISO 685A Dama LM3 >200,000 -10.72 0.031 -0.28 30.62 0.132 6.5
ISO 685B -10.88 0.036 -1.18 29.69 0.111 5.2
ISO 685C -10.90 0.050 -2.51 28.32 0.016 4.3
ISO 685D -11.15 0.045 -2.79 28.03 0.035 3.4
ISO 685E -11.27 0.030 -3.11 27.70 0.055 2.2
ISO 685F -11.08 0.033 -2.83 27.99 0.063 1.0
average: -11.00 28.72
ISO 656A Gazella LM3 ~200ka -13.19 0.026 0.80 31.73 0.048 5.9
ISO 656B -13.22 0.011 0.69 31.62 0.041 4.8
ISO 656C -13.34 0.029 0.78 31.71 0.047 3.4
ISO 656D -12.89 0.017 1.20 32.14 0.068 2.3
ISO 656E -12.29 0.049 2.07 33.04 0.073 1.1
average: -12.99 32.05
ISO 654A Gazella LM3 170-200ka -12.34 0.018 0.34 31.25 0.048 12.4
ISO 654B -12.14 0.030 0.43 31.35 0.059 11.1
ISO 654C -11.91 0.029 0.53 31.45 0.043 10.0
ISO 654D -11.58 0.038 0.57 31.50 0.105 8.9
ISO 654E -10.99 0.016 1.33 32.27 0.047 7.9
ISO 654F -10.64 0.022 1.06 32.00 0.015 6.8
ISO 654G -10.33 0.027 1.10 32.04 0.040 5.7
ISO 654H -9.98 0.039 1.08 32.02 0.021 4.6
ISO 654I -9.63 0.046 0.88 31.81 0.069 3.5
ISO 654J -9.20 0.019 0.84 31.77 0.061 2.3
ISO 654K -8.74 0.015 0.72 31.65 0.054 1.1
average: -10.68 31.74
ISO 667A Gazella UM3 ~170ka -13.15 0.024 -1.42 29.44 0.055 20.3
ISO 667B -13.38 0.046 -2.08 28.76 0.02 18.8
44
ISO 667C -13.39 0.026 -1.64 29.22 0.028 17.1
ISO 667D -13.55 0.031 -1.91 28.94 0.105 15.4
ISO 667E -13.7 0.042 -1.64 29.22 0.078 13.6
ISO 667F -13.68 0.02 -1.58 29.28 0.037 11.9
ISO 667G -13.66 0.032 -0.94 29.94 0.024 10.0
ISO 667H -13.58 0.022 -0.47 30.42 0.137 8.3
ISO 667I -13.48 0.033 -0.23 30.67 0.038 6.9
ISO 667J -13.45 0.021 0.17 31.08 0.056 5.5
ISO 667K -13.28 0.054 -0.24 30.66 0.067 4.2
ISO 667L -13.22 0.025 -0.37 30.53 0.1 2.8
ISO 667M -13.17 0.025 -0.53 30.36 0.057 1.1
average: -13.44 29.89
ISO 669B -13.52 0.018 0.78 31.71 0.097 10.9
ISO 669C -13.48 0.026 1.6 32.56 0.06 9.9
ISO 669D -13.42 0.044 1.75 32.71 0.052 8.5
ISO 669E -13.14 0.049 1.8 32.76 0.033 7.3
ISO 669F -13.06 0.019 1.83 32.79 0.113 6.0
ISO 669G -13.13 0.023 1.68 32.64 0.054 4.7
ISO 669H -12.88 0.038 1.27 32.22 0.073 3.5
ISO 669I -12.9 0.019 0.32 31.24 0.022 2.3
ISO 669J -12.90 0.019 -0.80 30.08 0.093 1.1
average: -13.211 32.07
ISO 678A Dama UP4 ~170ka -11.22 0.041 -1.31 29.55 0.050 10.4
ISO 678B -11.08 0.041 -1.79 29.06 0.056 9.6
ISO 678C -11.28 0.018 -2.04 28.81 0.019 8.3
ISO 678D -11.23 0.015 -2.38 28.45 0.021 7.4
ISO 678E -11.34 0.032 -2.08 28.76 0.052 6.5
ISO 678F -11.21 0.020 -2.14 28.70 0.050 5.7
ISO 678G -11.17 0.046 -2.27 28.56 0.020 4.4
ISO 678H -11.00 0.037 -2.24 28.60 0.041 3.5
ISO 678I -11.03 0.026 -2.60 28.23 0.060 2.4
ISO 678J -11.47 0.014 -2.85 27.96 0.054 1.5
ISO 678K -11.58 0.053 -2.75 28.07 0.047 0.7
average: -11.24 28.61
ISO 672A Dama LM3 ~170ka -11.10 0.048 0.91 31.84 0.036 13.7
ISO 672B -11.49 0.026 1.18 32.12 0.043 12.2
ISO 672C -10.83 0.012 1.07 32.00 0.055 10.8
ISO 672D -10.83 0.012 2.15 33.13 0.027 9.6
ISO 672E -10.48 0.013 1.47 32.42 0.080 8.5
ISO 672F -10.43 0.027 1.07 32.01 0.071 7.2
ISO 672G -10.17 0.025 1.02 31.96 0.021 6.0
ISO 672H -10.03 0.058 0.24 31.15 0.139 4.8
ISO 672I/ -9.97 0.025 -0.50 30.40 0.079 3.1
ISO 672J -9.70 0.018 -1.36 29.51 0.039 1.4
average: -10.50 31.65
45
ISO 683A Dama UM3 ~170ka -12.65 0.027 0.52 31.44 0.038 12.3
ISO 683B -12.67 0.012 -0.36 30.54 0.055 11.0
ISO 683C -12.46 0.023 -0.75 30.14 0.076 9.8
ISO 683D -12.42 0.021 -0.96 29.91 0.063 8.2
ISO 683E -12.31 0.033 -1.17 29.70 0.014 6.9
ISO 683F -12.04 0.014 -1.70 29.15 0.071 5.7
ISO 683G -11.74 0.033 -1.68 29.18 0.044 4.3
ISO 683H -11.88 0.013 -1.92 28.93 0.060 3.0
ISO 683I -11.32 0.013 -2.13 28.71 0.026 1.9
ISO 683J -10.76 0.019 -1.60 29.26 0.087 0.7
average: -12.03 29.70
ISO 653A Gazella LM3 ~170ka -13.50 0.064 -1.81 29.04 0.152 13.9
ISO 653B -13.42 0.050 -1.37 29.50 0.039 12.7
ISO 653C -13.39 0.055 -0.12 30.78 0.050 11.9
ISO 653D -13.33 0.012 0.60 31.53 0.056 10.8
ISO 653E -13.15 0.013 1.39 32.34 0.029 9.5
ISO 653F -13.06 0.019 0.63 31.56 0.031 8.3
ISO 653G -13.01 0.043 0.37 31.29 0.042 7.0
ISO 653H -12.75 0.014 0.30 31.21 0.060 5.9
ISO 653I -12.46 0.025 0.86 31.79 0.040 4.7
ISO 653J -12.23 0.021 0.48 31.40 0.037 3.3
ISO 653K -11.88 0.039 0.54 31.46 0.082 2.4
ISO 653L -11.69 0.018 -0.14 30.77 0.012 1.2
average: -12.82 31.05
ISO 663B -12.78 0.010 -0.26 30.64 0.062 18.1
ISO 663C -13.06 0.020 -0.38 30.51 0.031 16.6
ISO 663D -12.99 0.031 -0.65 30.24 0.034 15.1
ISO 663E -13.28 0.027 -0.23 30.67 0.031 13.7
ISO 663F -13.02 0.035 -0.34 30.56 0.070 12.2
ISO 663G -13.09 0.052 -0.22 30.68 0.085 10.8
ISO 663H -13.05 0.019 0.17 31.08 0.042 9.1
ISO 663I -12.78 0.011 0.58 31.50 0.062 7.9
ISO 663J -12.60 0.009 -0.02 30.89 0.032 6.5
ISO 663K -12.28 0.044 0.29 31.21 0.074 5.2
ISO 663L -11.27 0.027 0.04 30.95 0.068 4.0
ISO 663M -12.00 0.022 0.11 31.02 0.030 2.8
ISO 663N -11.69 0.018 0.14 31.05 0.073 1.3
average: -12.61 30.88
ISO 662A Gazella UM3 150-170ka -12.88 0.033 -1.01 29.86 0.058 8.6
ISO 662B -13.50 0.017 -0.57 30.31 0.061 7.4
ISO 662C -13.41 0.017 -0.23 30.67 0.049 6.0
ISO 662D -12.99 0.022 -0.35 30.54 0.058 4.6
ISO 662E -13.08 0.017 -0.08 30.82 0.073 3.5
ISO 662F -13.05 0.021 -0.62 30.26 0.061 2.1
46
ISO 662G -12.23 0.032 0.68 31.60 0.062 1.2
average: -13.02 30.58
ISO 659A Gazella LM3 150-170ka -13.50 0.049 -2.47 28.36 0.082 11.0
ISO 659B -13.63 0.023 -2.49 28.34 0.070 9.8
ISO 659C -13.90 0.036 -2.65 28.17 0.055 7.9
ISO 659D -13.89 0.035 -2.51 28.32 0.085 6.5
ISO 659E -13.93 0.014 -2.35 28.48 0.037 5.0
ISO 659F -13.62 0.012 -1.93 28.92 0.059 3.7
ISO 659G -13.57 0.018 -1.42 29.44 0.035 2.2
ISO 659H -13.81 0.037 -1.33 29.54 0.049 0.9
average: -13.73 28.70
ISO 650B -13.49 0.024 1.05 31.99 0.015 15.0
ISO 650C -13.63 0.023 -0.31 30.59 0.036 13.8
ISO 650D -13.92 0.025 -0.55 30.34 0.030 12.5
ISO 650E -13.94 0.037 -0.69 30.20 0.067 11.2
ISO 650F -14.04 0.030 -0.19 30.72 0.036 9.9
ISO 650G -14.15 0.015 -0.05 30.86 0.032 8.6
ISO 650H -14.22 0.023 0.50 31.43 0.096 7.3
ISO 650I -14.08 0.050 -0.04 30.87 0.041 6.0
ISO 650J -13.90 0.043 0.19 31.10 0.059 4.7
ISO 650K -13.70 0.020 -0.09 30.82 0.053 3.5
ISO 650L -13.42 0.018 0.63 31.56 0.045 2.2
ISO 650M -13.53 0.009 1.00 31.93 0.050 1.1
average: -13.78 31.17
ISO 668A Gazella LM3 70-100ka -12.45 0.061 -0.89 29.99 0.124 16.8
ISO 668B -12.92 0.037 -1.39 29.47 0.055 15.3
ISO 668C -13.60 0.048 -0.85 30.03 0.077 14.0
ISO 668D -13.60 0.022 -1.29 29.58 0.047 12.8
ISO 668E -13.31 0.028 -0.88 30.00 0.038 11.5
ISO 668F -12.95 0.044 -1.79 29.06 0.057 10.6
ISO 668G -13.27 0.022 -0.98 29.90 0.055 9.3
ISO 668H -12.95 0.058 -0.85 30.03 0.113 8.1
ISO 668I -12.98 0.025 -0.47 30.42 0.065 6.8
ISO 668J -12.73 0.053 -0.02 30.89 0.038 5.4
ISO 668K -12.29 0.067 -0.05 30.86 0.076 4.0
ISO 668L -11.75 0.033 0.28 31.20 0.112 2.9
ISO 668M -11.31 0.056 0.15 31.06 0.048 1.3
average: -12.78 30.19
ISO 666A Gazella UM3 70-100ka -11.64 0.030 1.82 32.79 0.053 15.7
ISO 666B -11.88 0.008 2.32 33.30 0.044 14.1
ISO 666C -11.95 0.020 2.25 33.22 0.046 12.8
ISO 666D -12.22 0.013 3.24 34.25 0.053 11.5
ISO 666E -12.66 0.037 2.17 33.15 0.085 10.3
ISO 666F -12.86 0.065 2.06 33.03 0.025 9.2
47
ISO 662G -13.12 0.019 1.26 32.21 0.029 7.9
ISO 666H -13.41 0.009 0.79 31.72 0.034 6.6
ISO 666I -13.37 0.029 -0.26 30.64 0.093 5.1
ISO 666J -13.42 0.044 -0.40 30.49 0.019 3.9
ISO 666K -13.08 0.020 -0.49 30.40 0.021 2.5
ISO 666L -13.04 0.015 0.05 30.96 0.060 1.1
average: -12.72 32.18
HAYONIM-AURIGNACIAN
26-28ka
ISO 642A Dama LP4 uncal. -11.29 0.034 2.23 33.21 0.042 8.9
ISO 642B -10.95 0.020 0.89 31.82 0.060 7.8
ISO 642C -10.62 0.101 -0.28 30.62 0.051 6.7
ISO 642D -10.12 0.086 -1.19 29.68 0.199 5.6
ISO 642E -10.41 0.160 -1.11 29.76 0.075 4.5
ISO 642F -10.28 0.020 -0.90 29.98 0.107 3.2
ISO 642G -10.23 0.028 -1.20 29.67 0.078 2.0
ISO 642H -10.08 0.101 -1.07 29.80 0.102 1.1
average: -10.50 30.57
ISO 633A Gazella UM3 -13.56 0.045 0.55 31.47 0.039 11.2
ISO 633B -13.48 0.037 0.36 31.28 0.083 10.0
ISO 633C -13.43 0.042 0.80 31.73 0.085 8.8
ISO 633D -13.20 0.030 1.17 32.11 0.082 7.6
ISO 633E -13.14 0.024 1.44 32.40 0.068 6.7
ISO 633F -12.96 0.018 1.50 32.46 0.011 5.5
ISO 633G -12.86 0.033 1.88 32.84 0.029 4.6
ISO 633H -12.56 0.019 1.99 32.95 0.019 3.4
ISO 633I -12.61 0.024 1.51 32.46 0.048 2.3
ISO 633J -12.54 0.039 1.77 32.73 0.070 1.1
average: -13.03 32.24
ISO 637A Gazella UM3 -13.12 0.026 0.61 31.54 0.068 8.3
ISO 637B -13.08 0.013 0.91 31.84 0.056 7.1
ISO 637C -12.95 0.021 1.53 32.48 0.070 6.0
ISO 637D -12.85 0.013 1.74 32.70 0.032 4.9
ISO 637E -12.69 0.037 2.25 33.23 0.052 3.8
ISO 637F -12.68 0.051 2.85 33.85 0.059 2.7
ISO 637G -12.64 0.015 3.57 34.59 0.027 1.5
ISO 637H -12.51 0.024 3.91 34.94 0.072 0.6
average: -12.82 33.15
ISO 639A Gazella UM3 -13.23 0.031 1.30 32.25 0.063 6.7
ISO 639B -13.03 0.045 1.59 32.55 0.025 5.8
ISO 639C -13.05 0.035 1.77 32.73 0.068 4.6
ISO 639D -12.74 0.033 1.40 32.35 0.067 3.5
ISO 639E -12.73 0.041 2.10 33.07 0.040 2.6
ISO 639F -12.42 0.037 2.18 33.16 0.029 1.4
ISO 639G -12.32 0.045 2.16 33.13 0.084 0.5
48
average: -12.79 32.75
ISO 638A Gazella UM3 -13.13 0.021 0.92 31.85 0.058 14.0
ISO 638B -13.11 0.021 0.84 31.77 0.051 12.8
ISO 638C -12.99 0.020 0.75 31.68 0.049 11.7
ISO 638D -12.96 0.022 1.59 32.55 0.071 10.6
ISO 638E -12.99 0.040 1.36 32.31 0.066 9.5
ISO 638F -12.99 0.014 1.28 32.22 0.028 8.4
ISO 638G -12.94 0.030 1.16 32.10 0.055 7.4
ISO 638H -12.99 0.017 0.73 31.66 0.043 6.3
ISO 638I -12.85 0.037 0.14 31.05 0.057 5.1
ISO 638J -12.79 0.054 -0.07 30.83 0.025 4.0
ISO 638K -12.58 0.040 -0.92 29.96 0.029 2.7
ISO 638L -12.34 0.027 -1.99 28.86 0.110 1.9
ISO 638M -12.39 0.039 -2.28 28.56 0.050 0.8
average: -12.85 31.18
ISO 640A Dama LP4 -11.40 0.024 1.03 31.97 0.046 10.9
ISO 640B -11.41 0.007 0.34 31.26 0.083 9.7
ISO 640C -11.45 0.022 0.42 31.34 0.050 8.5
ISO 640D -11.22 0.035 -0.15 30.75 0.054 7.8
ISO 640E -11.14 0.035 -0.16 30.74 0.037 6.6
ISO 640F -11.03 0.030 0.05 30.96 0.041 5.4
ISO 640G -11.25 0.017 -0.02 30.88 0.041 4.6
ISO 640H -11.62 0.034 0.76 31.69 0.039 3.9
ISO 640I -11.09 0.032 0.96 31.89 0.053 2.9
ISO 640J -11.38 0.031 1.11 32.06 0.058 1.7
ISO 640K -11.64 0.020 1.68 32.64 0.021 0.6
average: -11.33 31.47
ISO 641A Dama LM3 -10.35 0.022 0.60 31.53 0.054 10.3
ISO 641B -10.05 0.017 0.37 31.29 0.042 9.1
ISO 641C -10.10 0.028 0.03 30.93 0.125 8.4
ISO 641D -10.05 0.036 0.03 30.94 0.026 7.2
ISO 641E -9.94 0.029 -0.76 30.12 0.053 6.1
ISO 641F -10.18 0.032 -1.25 29.62 0.063 5.0
ISO 641G -10.21 0.008 -1.35 29.51 0.031 4.1
ISO 641H -9.96 0.015 -0.85 30.03 0.054 3.1
ISO 641I -10.27 0.015 -1.46 29.40 0.057 2.0
ISO 641J -10.54 0.012 -1.78 29.07 0.049 0.9
average: -10.17 30.24
ISO 645A Dama UP4 . -11.76 0.020 0.62 31.54 0.022 11.6
ISO 645B -11.66 0.017 0.28 31.20 0.076 10.4
ISO 645C -11.36 0.017 -0.29 30.61 0.065 9.5
ISO 645D -11.27 0.010 -0.29 30.60 0.042 8.2
ISO 645E -11.37 0.055 -0.28 30.62 0.089 7.0
ISO 645F -11.63 0.023 -0.35 30.55 0.037 5.9
ISO 645G -11.54 0.013 -0.24 30.66 0.072 4.8
49
ISO 645H -11.76 0.052 0.04 30.94 0.034 3.6
ISO 645I -11.76 0.019 0.26 31.17 0.058 2.4
ISO 645J -11.79 0.016 0.33 31.24 0.048 1.2
average: -11.59 30.91
MEGED
18-19ka
ISO 631A Dama LM3 uncal. -11.15 0.013 -0.25 30.65 0.033 16.0
ISO 631B -11.49 0.033 0.66 31.59 0.051 14.4
ISO 631C -11.68 0.022 0.63 31.56 0.041 12.9
ISO 631D -11.50 0.029 -0.01 30.90 0.018 11.3
ISO 631E -11.28 0.029 0.23 31.14 0.019 10.0
ISO 631F -10.59 0.015 0.27 31.19 0.029 8.2
ISO 631G -10.65 0.012 0.44 31.36 0.058 6.9
ISO 631H -10.47 0.038 0.56 31.48 0.046 5.6
ISO 631I -10.60 0.037 -0.33 30.57 0.023 4.4
ISO 631J -10.40 0.009 -1.17 29.70 0.095 3.1
ISO 631K -10.41 0.016 -1.15 29.72 0.074 1.8
ISO 631L -10.33 0.013 -1.71 29.15 0.036 0.6
average: -10.88 30.75
ISO 620A Gazella LM3 -11.92 0.024 -0.56 30.33 0.015 18.2
ISO 620B -12.05 0.017 -0.25 30.65 0.045 16.8
ISO 620C -12.50 0.052 -0.32 30.57 0.096 15.0
ISO 620D -12.68 0.040 0.35 31.27 0.108 13.4
ISO 620E -12.91 0.029 0.02 30.93 0.034 11.8
ISO 620F -12.44 0.027 0.60 31.53 0.034 9.5
ISO 620G -12.19 0.016 1.02 31.96 0.047 7.6
ISO 620H -12.19 0.014 0.15 31.06 0.016 6.3
ISO 620I -12.22 0.031 0.88 31.82 0.055 4.7
ISO 620J -12.19 0.015 0.76 31.69 0.036 3.1
ISO 620K -12.10 0.031 1.79 32.75 0.030 1.4
average: -12.31 31.32
ISO 619A Gazella UM3 -12.41 0.026 -0.29 30.61 0.035 14.1
ISO 619B -12.56 0.042 -0.44 30.45 0.039 13.0
ISO 619C -12.39 0.028 0.82 31.76 0.065 11.8
ISO 619D -12.38 0.036 1.11 32.05 0.028 10.4
ISO 619E -12.76 0.015 1.26 32.20 0.079 9.3
ISO 619F -12.29 0.019 0.66 31.59 0.018 8.4
ISO 619G -11.94 0.010 0.38 31.30 0.037 7.3
ISO 619H -12.33 0.028 1.21 32.15 0.077 6.1
ISO 619I -11.99 0.047 1.52 32.47 0.047 5.2
ISO 619J -12.24 0.024 1.20 32.15 0.057 4.0
ISO 619K -12.27 0.030 1.50 32.45 0.027 2.9
ISO 619L -12.49 0.012 2.37 33.35 0.098 1.8
ISO 619M -12.36 0.021 2.47 33.45 0.063 0.7
average: -12.34 32.00
50
ISO 628A Dama LM3 -10.85 0.038 -0.61 30.28 0.034 7.8
ISO 628B -11.24 0.025 -0.31 30.59 0.050 6.3
ISO 628C -11.19 0.034 -0.91 29.97 0.066 5.0
ISO 628D -11.19 0.037 -1.12 29.75 0.036 3.9
ISO 628E -11.71 0.042 -2.39 28.44 0.049 2.5
ISO 628F -11.73 0.034 -2.31 28.53 0.053 1.6
ISO 628G -11.77 0.022 -1.83 29.02 0.067 0.7
average: -11.38 29.51
HAYONIM-KEBARAN
14-17ka
ISO 609A Gazella UM3 uncal. -13.14 0.028 -0.16 30.74 0.061 13.0
ISO 609B -12.95 0.028 -0.04 30.86 0.051 11.5
ISO 609C -13.11 0.013 0.19 31.10 0.047 10.3
ISO 609D -13.18 0.043 0.20 31.12 0.049 9.2
ISO 609E -13.00 0.010 -0.19 30.71 0.065 8.1
ISO 609F -12.98 0.017 0.01 30.92 0.084 7.0
ISO 609G -12.93 0.023 0.00 30.91 0.055 5.8
ISO 609H -12.95 0.049 -0.16 30.74 0.091 4.9
ISO 609I -13.00 0.028 -0.12 30.79 0.022 3.8
ISO 609J -12.51 0.050 0.03 30.94 0.043 2.6
ISO 609K -12.44 0.029 0.41 31.33 0.065 1.2
average: -12.93 30.92
ISO 608A Gazella UM3 -13.96 0.030 -0.62 30.27 0.075 17.2
ISO 608B -13.96 0.021 -0.08 30.82 0.061 16.0
ISO 608C -13.51 0.029 -0.54 30.35 0.027 14.9
ISO 608D -13.61 0.022 -0.17 30.73 0.026 13.8
ISO 608E -13.47 0.023 -0.47 30.42 0.060 12.4
ISO 608F -13.30 0.025 -0.33 30.56 0.041 11.2
ISO 608G -12.76 0.011 0.37 31.29 0.060 10.1
ISO 608H -13.35 0.058 0.28 31.20 0.106 9.0
ISO 608I -12.84 0.036 0.27 31.19 0.035 7.9
ISO 608J -13.00 0.028 0.54 31.46 0.056 6.6
ISO 608K -12.50 0.028 0.40 31.32 0.094 5.4
ISO 608L -12.31 0.036 0.55 31.47 0.045 4.5
ISO 608M -12.36 0.017 1.12 32.06 0.023 3.3
ISO 608N -12.34 0.034 0.50 31.42 0.073 2.1
ISO 608O -11.82 0.039 0.51 31.43 0.053 1.0
average: -13.01 31.07
ISO 613A Dama LM3 -12.02 0.064 -0.25 30.64 0.069 13.7
ISO 613B -11.78 0.029 0.22 31.14 0.067 12.8
ISO 613C -12.17 0.020 -0.13 30.78 0.104 11.5
ISO 613D -11.91 0.034 -0.15 30.75 0.076 10.3
ISO 613E -11.19 0.162 -0.14 30.76 0.245 9.0
ISO 613F -11.44 0.032 -0.34 30.56 0.058 7.5
ISO 613G -11.24 0.111 -1.27 29.59 0.045 6.2
51
ISO 613H -11.15 0.017 -1.57 29.29 0.036 5.0
ISO 613I -10.99 0.042 -1.87 28.98 0.036 3.9
ISO 613J -11.17 0.031 -1.72 29.13 0.075 2.8
ISO 613K -10.90 0.029 -1.85 29.00 0.020 1.9
ISO 613L -10.94 0.016 -1.99 28.86 0.030 0.7
average: -11.41 29.96
ISO 616A Dama LM3 -11.98 0.018 -0.04 30.86 0.031 13.1
ISO 616B -11.72 0.025 -0.72 30.16 0.041 11.9
ISO 616C -11.80 0.020 -1.28 29.58 0.068 10.7
ISO 616D -11.41 0.021 -2.54 28.28 0.071 9.0
ISO 616E -11.78 0.022 -2.46 28.37 0.028 7.8
ISO 616F -11.52 0.010 -2.49 28.34 0.046 6.5
ISO 616G -11.96 0.013 -2.09 28.75 0.107 5.2
ISO 616H -11.81 0.040 -2.15 28.69 0.028 4.0
ISO 616I -12.00 0.030 -1.92 28.93 0.067 2.9
ISO 616J -11.81 0.024 -1.92 28.93 0.042 1.8
ISO 616K -11.94 0.031 -1.76 29.09 0.035 0.6
average: -11.79 29.09
ISO 615A Dama UP4 -10.29 0.056 -0.09 30.82 0.087 7.9
ISO 615B -10.67 0.027 -0.22 30.68 0.024 6.5
ISO 615C -10.93 0.038 -0.06 30.85 0.080 5.6
ISO 615D -10.81 0.042 -1.58 29.28 0.077 4.3
ISO 615E -10.69 0.013 -1.78 29.07 0.046 3.1
ISO 615F -10.42 0.041 -2.02 28.82 0.038 2.2
ISO 615G -10.34 0.063 -1.88 28.97 0.133 1.0
average: -10.59 29.78
HAYONIM-NATUFIAN
11-13ka
ISO 605A Gazella LM2 uncal. -12.20 0.033 -3.38 27.43 0.046 13.3
ISO 605B -12.30 0.033 -3.10 27.71 0.074 12.9
ISO 605C -12.45 0.017 -3.03 27.78 0.027 11.6
ISO 605D -12.44 0.025 -3.14 27.67 0.025 10.4
ISO 605E -12.60 0.030 -2.90 27.92 0.027 9.1
ISO 605F -12.63 0.063 -2.89 27.93 0.032 7.9
ISO 605G -12.98 0.020 -2.54 28.29 0.063 6.8
ISO 605H -13.34 0.042 -2.98 27.83 0.020 5.4
ISO 605I -13.57 0.019 -2.33 28.50 0.025 4.0
ISO 605J -13.72 0.028 -2.51 28.32 0.058 2.9
ISO 605K -13.47 0.020 -2.84 27.98 0.086 1.6
average: -12.88 27.94
ISO 604A Gazella LM2 -12.82 0.034 0.03 30.94 0.019 14.8
ISO 604B -12.92 0.017 -0.78 30.10 0.033 13.5
ISO 604C -12.77 0.061 -1.14 29.73 0.024 12.2
ISO 604D -12.88 0.022 -0.28 30.62 0.040 11.0
52
ISO 604E -13.02 0.030 -0.26 30.64 0.030 9.8
ISO 604F -13.14 0.028 -0.25 30.65 0.051 8.7
ISO 604G -13.07 0.037 -0.53 30.36 0.057 7.6
ISO 604H -13.03 0.023 -0.51 30.38 0.060 6.5
ISO 604I -12.97 0.047 -0.41 30.48 0.053 5.5
ISO 604J -13.01 0.024 -0.12 30.79 0.030 4.4
ISO 604K -13.09 0.030 -0.47 30.43 0.061 3.3
ISO 604L -12.99 0.013 -0.30 30.60 0.027 2.2
ISO 604M -12.97 0.041 -0.22 30.68 0.052 1.0
average: -12.98 30.49
ISO 602A Gazella LP4 -13.19 0.023 2.46 33.44 0.066 7.6
ISO 602B -13.11 0.014 2.67 33.66 0.040 6.4
ISO 602C -13.00 0.012 2.68 33.67 0.060 5.3
ISO 602D -12.54 0.036 2.67 33.66 0.056 4.4
ISO 602E -12.74 0.023 2.79 33.78 0.052 3.2
ISO 602F -12.57 0.034 2.60 33.59 0.046 2.1
ISO 602G -12.88 0.056 2.60 33.58 0.076 0.7
average: -12.86 33.63
ISO 692A Gazella UM3 -13.12 0.034 -1.53 29.33 0.117 6.3
ISO 692B -13.38 0.029 -1.17 29.70 0.058 5.0
ISO 692C -12.82 0.042 -0.63 30.26 0.076 4.1
ISO 692D -13.26 0.011 -0.24 30.66 0.026 3.1
ISO 692E -13.47 0.035 -0.62 30.27 0.079 2.0
ISO 692F -13.75 0.012 -1.11 29.77 0.051 0.9
average: -13.30 30.00
ISO 689A Gazella LM2 -13.45 0.012 1.09 32.03 0.093 9.6
ISO 689B -13.82 0.035 1.18 32.12 0.056 8.5
ISO 689C -13.61 0.017 0.81 31.74 0.013 7.6
ISO 689D -13.65 0.045 0.07 30.98 0.052 6.5
ISO 689E -13.37 0.029 -0.56 30.33 0.028 5.7
ISO 689F -13.32 0.043 -1.64 29.22 0.102 4.9
ISO 689G -12.84 0.017 -1.88 28.97 0.068 3.7
ISO 689H -12.55 0.028 -2.03 28.82 0.047 2.8
ISO 689I -12.45 0.036 -2.11 28.74 0.064 1.9
ISO 689J -12.59 0.013 -2.01 28.83 0.070 1.0
average: -13.16 30.18
ISO 601A Gazella UP4 -12.11 0.044 0.62 31.55 0.106 4.7
ISO 601B -11.87 0.023 0.50 31.42 0.052 3.8
ISO 601C -12.34 0.019 0.63 31.55 0.077 2.6
ISO 601D -12.51 0.024 0.84 31.77 0.050 1.7
ISO 601E -12.52 0.017 0.99 31.93 0.032 0.8
average: -12.27 31.64
53
HOLOCENE
4000-
HOL 689A Bos M3 3000BP -8.26 0.037 -0.61 30.28 0.047 47.9
HOL 689B -8.20 0.015 -0.28 30.62 0.074 45.4
HOL 689C -8.44 0.025 -1.91 28.93 0.053 43.0
HOL 689D -8.63 0.020 -2.80 28.02 0.076 40.9
HOL 689E -8.61 0.037 -2.91 27.90 0.027 39.0
HOL 689F -9.29 0.017 -2.78 28.04 0.024 37.0
HOL 689G -9.71 0.022 -3.24 27.57 0.043 35.1
HOL 689H -10.12 0.018 -2.73 28.09 0.063 33.5
HOL 689I -10.35 0.031 -3.45 27.35 0.029 31.8
HOL 689J -10.06 0.027 -2.31 28.53 0.033 30.0
HOL 689K -10.17 0.026 -2.48 28.35 0.054 28.6
HOL 689L -9.67 0.023 -1.88 28.97 0.056 27.2
HOL 689M -9.96 0.019 -1.58 29.28 0.111 25.5
HOL 689N -10.43 0.032 -1.72 29.13 0.026 23.7
HOL 689O -10.53 0.048 -1.13 29.74 0.047 21.9
HOL 689P -10.68 0.037 -0.92 29.96 0.033 20.5
HOL 689Q -10.47 0.024 -1.12 29.75 0.048 18.9
HOL 689R -10.79 0.033 -1.18 29.69 0.050 17.0
HOL 689S -10.56 0.020 -1.02 29.85 0.091 14.8
HOL 689T -10.27 0.030 -0.86 30.02 0.052 13.0
HOL 689U -10.48 0.028 -1.17 29.70 0.022 10.9
HOL 689V -10.08 0.026 -0.29 30.61 0.042 8.6
HOL 689W -9.94 0.014 -0.98 29.90 0.068 6.5
HOL 689X -9.60 0.039 -0.43 30.46 0.080 4.4
HOL 689Y -9.41 0.013 -1.09 29.79 0.054 3.0
HOL 689Z -8.78 0.023 -1.04 29.84 0.018 1.7
average: -9.75 29.24
4000-
HOL 753A Capra M3 3000BP -8.33 0.033 -0.20 30.70 0.045 19.6
HOL 753B -7.75 0.008 -1.27 29.60 0.080 18.1
HOL 753C -7.35 0.023 -1.35 29.51 0.051 16.4
HOL 753D -7.38 0.030 -2.07 28.77 0.059 14.5
HOL 753E -7.30 0.034 -1.55 29.31 0.035 13.2
HOL 753F -7.61 0.021 -1.73 29.12 0.050 11.9
HOL 753G -7.80 0.033 -1.81 29.04 0.050 10.8
HOL 753H -7.64 0.024 -1.30 29.56 0.027 9.3
HOL 753I -7.67 0.031 -1.36 29.50 0.031 8.0
HOL 753J -7.55 0.032 -2.03 28.82 0.034 6.9
HOL 753K -7.55 0.016 -0.48 30.41 0.028 5.4
HOL 753L -8.56 0.036 1.56 32.52 0.056 4.1
HOL 753M -9.73 0.028 0.88 31.81 0.033 2.9
HOL 753N -8.42 0.019 1.73 32.69 0.047 1.3
average: -7.90 30.10
4000-
HOL 370A Capra M3 3000BP -9.28 0.030 0.27 31.18 0.049 24.7
HOL 370B -9.42 0.016 -0.16 30.74 0.042 23.4
54
HOL 370C -9.52 0.033 -0.15 30.75 0.060 21.9
HOL 370D -9.75 0.012 -0.63 30.26 0.039 20.6
HOL 370E -9.94 0.049 -0.25 30.65 0.050 19.0
HOL 370F -10.15 0.012 -0.09 30.81 0.035 17.4
HOL 370G -10.32 0.017 -0.24 30.66 0.050 16.0
HOL 370H -9.75 0.017 0.78 31.72 0.066 14.5
HOL 370I -9.62 0.018 0.61 31.53 0.031 13.3
HOL 370J -9.43 0.042 1.36 32.30 0.059 11.9
HOL 370K -9.35 0.031 1.60 32.55 0.043 10.4
HOL 370L -8.91 0.023 1.69 32.65 0.085 9.1
HOL 370M -8.63 0.016 1.42 32.37 0.049 7.3
HOL 370N -8.55 0.014 0.84 31.77 0.098 6.1
HOL 370O -8.82 0.028 0.41 31.33 0.044 5.0
HOL 370P -8.66 0.029 -0.69 30.19 0.030 3.8
HOL 370Q -9.11 0.023 -0.76 30.13 0.023 2.5
HOL 370R -9.18 0.043 -0.79 30.09 0.056 1.2
average: -9.35 31.20
HOL 154A Bos M3 ~3000BP -9.11 0.041 0.06 30.96 0.039 43.2
HOL 154B -8.66 0.041 -1.15 29.72 0.042 41.8
HOL 154C -8.58 0.041 0.11 31.02 0.046 40.5
HOL 154D -9.08 0.024 -0.46 30.44 0.066 39.2
HOL 154E -8.87 0.051 -0.57 30.32 0.073 38.1
HOL 154F -9.09 0.019 -1.17 29.70 0.042 37.0
HOL 154G -9.03 0.047 -0.08 30.82 0.057 35.5
HOL 154H -9.31 0.023 -1.14 29.73 0.039 34.0
HOL 154I -9.23 0.023 -1.15 29.72 0.051 32.8
HOL 154J -9.20 0.022 -1.25 29.61 0.057 31.2
HOL 154K -9.44 0.046 -1.71 29.14 0.023 29.9
HOL 154L -9.63 0.021 -1.57 29.29 0.051 28.5
HOL 154M -9.36 0.012 -1.73 29.12 0.024 27.0
HOL 154N -9.32 0.022 -1.86 28.99 0.054 25.6
HOL 154O -9.60 0.040 -1.86 28.99 0.062 24.1
HOL 154P -9.84 0.028 -1.55 29.31 0.037 22.4
HOL 154Q -10.15 0.025 -1.23 29.64 0.052 20.9
HOL 154R -10.34 0.014 -1.43 29.44 0.045 19.0
HOL 154S -10.20 0.034 -1.68 29.17 0.049 16.7
HOL 154T -10.16 0.015 -1.63 29.22 0.024 15.0
HOL 154U -9.79 0.010 -1.93 28.91 0.032 13.1
HOL 154V -9.12 0.026 -2.22 28.62 0.061 11.8
HOL 154W -9.18 0.013 -2.38 28.45 0.074 10.0
HOL 154X -9.22 0.036 -2.98 27.84 0.067 7.9
HOL 154Y -9.58 0.041 -2.86 27.96 0.029 5.8
HOL 154Z -9.06 0.025 -3.89 26.90 0.054 3.9
HOL 154AA -8.49 0.030 -2.95 27.87 0.021 2.5
HOL 154BB -8.03 0.026 -2.72 28.10 0.036 1.2
average: -9.31 29.25
55
Appendix 2
Intra-tooth Microsampling Plots; δ13C and δ18O Covariance Plots
QC 411 Covariance QC 411
30.00 -10.20
-10.40

29.50 -10.60
R2 = 0.3716
-10.80
29.00 -11.00
-11.20
28.50 -11.40
-11.60
28.00 -11.80
-12.00
27.50
27.50 28.00 28.50 29.00 29.50 30.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
QC 411
-10.20
-10.40
-10.60
-10.80
-11.00
-11.20
-11.40
-11.60
-11.80
-12.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
30.00 -9.8
-10
29.50 R2 = 0.2321
-10.2
29.00
-10.4
28.50 -10.6
28.00 -10.8
-11
27.50
27.5 28 28.5 29 29.5 30
0.0 2.0 4.0 6.0 8.0 10.0 12.0
QC 417
-9.80
-10.00
-10.20
-10.40
-10.60
-10.80
-11.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
56
28.80 -11.00
-11.10

28.60
δ18O bioapatite (‰) R2 = 0.1484
28.40 -11.20
-11.30
28.20
-11.40
28.00
-11.50
27.80
-11.60
27.60 27.60 27.80 28.00 28.20 28.40 28.60 28.80
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 δ18O bioapatite (‰)
QC 425
-10.00
-10.40
-10.80
-11.20
-11.60
-12.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0
33.00 -10.60
-10.80
32.00
R2 = 0.0951
-11.00
31.00
-11.20
30.00
-11.40
29.00 -11.60
-11.80
28.00
28.00 29.00 30.00 31.00 32.00 33.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
QC 430
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
57
30.00 -10.40
-10.60
29.50

R2 = 0.1047
δ18O bioapatite (‰) -10.80
29.00 -11.00
-11.20
28.50
-11.40
28.00 -11.60
27.50 -11.80
-12.00
27.00 27.00 27.50 28.00 28.50 29.00 29.50 30.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
QC 426
-10.40
-10.60
-10.80
-11.00
-11.20
-11.40
-11.60
-11.80
-12.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
30.50 -10.70
30.00 -10.80
29.50 -10.90
29.00 -11.00
R2 = 0.1294
28.50 -11.10
28.00 -11.20
27.50 -11.30
27.00 27.00 27.50 28.00 28.50 29.00 29.50 30.00 30.50
0.0 2.0 4.0 6.0 8.0 10.0 δ18O bioapatite (‰)
QC 432
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 2.0 4.0 6.0 8.0 10.0
58
29.50 -10.40
-10.50
29.00

28.50
-10.70
28.00 R2 = 0.1419
-10.80
27.50
-10.90
27.00
-11.00
26.50
-11.10
26.00 26.00 26.50 27.00 27.50 28.00 28.50 29.00 29.50
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 δ18O bioapatite (‰)
QC 428
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0
32.00 -11.20
31.50 -11.40
31.00 -11.60
30.50 R2 = 0.9854
-11.80
30.00
-12.00
29.50
-12.20
29.00
-12.40
28.50 28.50 29.00 29.50 30.00 30.50 31.00 31.50 32.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 δ18O bioapatite (‰)
QC 438
-11.20
-11.40
-11.60
-11.80
-12.00
-12.20
-12.40
0.0 2.0 4.0 6.0 8.0 10.0 12.0
59
32.00 -10.10
-10.20
31.00

-10.30
30.00 -10.40
R2 = 0.0835
-10.50
29.00
-10.60
28.00 -10.70
-10.80
27.00
28.50 29.00 29.50 30.00 30.50 31.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
QC 442
-10.00
-10.40
-10.80
-11.20
-11.60
-12.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
ISO 681 Covariance ISO 681
31.50 -10.20
31.00
-10.40
30.50
-10.60
30.00
R2 = 0.0008
29.50 -10.80
29.00 -11.00
28.50
-11.20
28.00 28.00 28.50 29.00 29.50 30.00 30.50 31.00 31.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 δ18O bioapatite (‰)
ISO 681
-10.20
-10.40
-10.60
-10.80
-11.00
-11.20
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
60
31.00 -10.20
30.50

-10.40
30.00 R2 = 0.0008
-10.60
29.50
-10.80
29.00
28.50 -11.00
28.00 -11.20
27.50 28.00 28.50 29.00 29.50 30.00 30.50 31.00 31.50
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 δ18O bioapatite (‰)
ISO 685
-10.60
-10.70
-10.80
-10.90
-11.00
-11.10
-11.20
-11.30
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0
34.00 -12.20
R2 = 0.9704
33.50 -12.40
33.00 -12.60
-12.80
32.50
-13.00
32.00
-13.20
31.50
-13.40
31.00 31.50 32.00 32.50 33.00 33.50
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 δ18O bioapatite (‰)
ISO 656
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0
61
33.00 -8.00

-9.00
δ18O bioapatite (‰) 32.50
-10.00
32.00 -11.00
R2 = 0.264
-12.00
31.50
-13.00
31.00 31.20 31.40 31.60 31.80 32.00 32.20 32.40
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 δ18O bioapatite (‰)
ISO 654
-8.00
-9.00
-10.00
-11.00
-12.00
-13.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
31.50 -13.00
-13.10
31.00
-13.20
30.50 -13.30
R2 = 0.0656
-13.40
30.00
-13.50
29.50 -13.60
-13.70
29.00
-13.80
28.50 28.50 29.00 29.50 30.00 30.50 31.00 31.50
0.0 5.0 10.0 15.0 20.0 25.0 δ18O bioapatite (‰)
ISO 667
-13.1
-13.2
-13.3
-13.4
-13.5
-13.6
-13.7
-13.8
0.0 5.0 10.0 15.0 20.0 25.0
62
33.00 -12.80
32.50

-13.00
32.00
-13.20
31.50 R2 = 0.0997
31.00 -13.40
30.50 -13.60
30.00
-13.80
29.50 29.50 30.00 30.50 31.00 31.50 32.00 32.50 33.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 δ18O bioapatite (‰)
ISO 669
-12.80
-13.00
-13.20
-13.40
-13.60
-13.80
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
33.00 -10.80
32.50
-11.00
32.00
-11.20
31.50 R2 = 0.1524
31.00 -11.40
30.50 -11.60
30.00
-11.80
29.50 27.50 28.00 28.50 29.00 29.50 30.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 δ18O bioapatite (‰)
ISO 678
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0
63
33.50 -9.40
33.00
-9.80

δ18O bioapatite (‰) 32.50
32.00 -10.20
R2 = 0.4859
31.50 -10.60
31.00
-11.00
30.50
30.00 -11.40
29.50 -11.80
29.00 29.00 30.00 31.00 32.00 33.00 34.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
ISO 672
-9.00
-9.50
-10.00
-10.50
-11.00
-11.50
-12.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
32.00 -10.50
31.50 -11.00
31.00 -11.50
30.50 -12.00
R2 = 0.5351
30.00 -12.50
29.50
-13.00
29.00
-13.50
28.50 28.50 29.00 29.50 30.00 30.50 31.00 31.50 32.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 δ18O bioapatite (‰)
ISO 683
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
64
33.00 -11.00
-11.50

32.00
-12.00
31.00 R2 = 0.114
-12.50
30.00 -13.00
-13.50
29.00
-14.00
28.00 28.00 29.00 30.00 31.00 32.00 33.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 δ18O bioapatite (‰)
ISO 653
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
32.00 -11.00
-11.50
31.50
31.00 -12.00
R2 = 0.163
-12.50
30.50
-13.00
30.00
-13.50
29.50
-14.00
29.00 30.00 30.20 30.40 30.60 30.80 31.00 31.20 31.40 31.60
0.0 5.0 10.0 15.0 20.0 25.0 δ18O bioapatite (‰)
ISO 663
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
0.0 5.0 10.0 15.0 20.0 25.0
65
32.00 -12.00
31.50

-12.40
31.00
-12.80
30.50 R2 = 0.3429
-13.20
30.00
-13.60
29.50
-14.00
29.00 29.50 30.00 30.50 31.00 31.50 32.00
0.0 2.0 4.0 6.0 8.0 10.0
ISO 662
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
0.0 2.0 4.0 6.0 8.0 10.0
30.00 -13.20
29.50 -13.40
29.00
-13.60
28.50
-13.80
28.00 R2 = 0.0599
-14.00
27.50
-14.20
27.00
28.00 28.50 29.00 29.50 30.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
ISO 659
-12.00
-12.50
-13.00
-13.50
-14.00
-14.50
-15.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
66
34.00 -13.00
-13.20

33.00
32.00 -13.60
R2 = 0.4577
-13.80
31.00
-14.00
-14.20
30.00
-14.40
29.00 30.00 30.50 31.00 31.50 32.00 32.50 33.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0 δ18O bioapatite (‰)
ISO 650
-13.00
-13.20
-13.40
-13.60
-13.80
-14.00
-14.20
-14.40
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0
33.00 -11.00
-11.50
32.00
-12.00
31.00
-12.50
30.00 -13.00
R2 = 0.4968
29.00 -13.50
-14.00
28.00
28.50 29.00 29.50 30.00 30.50 31.00 31.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0
ISO 668
-10.00
-11.00
-12.00
-13.00
-14.00
-15.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0
67
35.00 -11.00
34.00 -11.50

33.00 -12.00
32.00 -12.50
-13.00
31.00 R2 = 0.5632
-13.50
30.00
-14.00
29.00 30.00 31.00 32.00 33.00 34.00 35.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0
ISO 666
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0
34.00 -9.80
33.00 -10.20
32.00 -10.60
R2 = 0.9363
31.00 -11.00
-11.40
30.00
-11.80
29.00
29.00 30.00 31.00 32.00 33.00 34.00
0.0 2.0 4.0 6.0 8.0 10.0
ISO 642
-10.00
-10.20
-10.40
-10.60
-10.80
-11.00
-11.20
-11.40
0.0 2.0 4.0 6.0 8.0 10.0
68
33.50 -12.00

33.00 -12.40
32.50 -12.80
-13.20
32.00
R2 = 0.8226
-13.60
31.50
-14.00
31.00 31.00 31.50 32.00 32.50 33.00 33.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0
ISO 633
-12.40
-12.60
-12.80
-13.00
-13.20
-13.40
-13.60
-13.80
0.0 2.0 4.0 6.0 8.0 10.0 12.0
36.00 -12.40
-12.60
35.00
34.00 -12.80
R2 = 0.9323
-13.00
33.00
-13.20
32.00
-13.40
31.00
31.00 32.00 33.00 34.00 35.00 36.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0
ISO 637
-12.40
-12.50
-12.60
-12.70
-12.80
-12.90
-13.00
-13.10
-13.20
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0
69
34.00 -12.20
33.50 -12.40

33.00 -12.60
R2 = 0.6156
-12.80
32.50
-13.00
32.00
-13.20
31.50
-13.40
31.00 32.20 32.40 32.60 32.80 33.00 33.20 33.40
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0
ISO 639
-12.20
-12.40
-12.60
-12.80
-13.00
-13.20
-13.40
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0
33.00 -12.20
-12.40
32.00
-12.60
31.00
-12.80
30.00 -13.00
R2 = 0.8804
29.00 -13.20
-13.40
28.00
28.00 29.00 30.00 31.00 32.00 33.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
ISO 638
-12.00
-12.20
-12.40
-12.60
-12.80
-13.00
-13.20
-13.40
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
70
33.00 -10.60
32.50 -10.80

32.00 R2 = 0.3765
-11.20
31.50 -11.40
31.00 -11.60
-11.80
30.50
-12.00
30.00 30.50 31.00 31.50 32.00 32.50 33.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
ISO 640
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
-9.80
32.00
31.50 -10.00
31.00 -10.20
30.50 R2 = 0.1237
-10.40
30.00
-10.60
29.50
29.00 -10.80
28.50 29.00 29.50 30.00 30.50 31.00 31.50 32.00
28.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0 δ18O bioapatite (‰)
ISO 641
-9.00
-9.50
-10.00
-10.50
-11.00
-11.50
-12.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
71
31.60 -11.00
31.40

-11.20
R2 = 0.1535
31.20
-11.40
31.00
-11.60
30.80
-11.80
30.60
-12.00
30.40 0.00 2.00 4.00 6.00 8.00 10.00 12.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
ISO 645
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
33.00 -10.00
32.00 -10.40
31.00 -10.80
R2 = 0.362
30.00
-11.20
29.00
-11.60
28.00
-12.00
27.00 29.00 29.50 30.00 30.50 31.00 31.50 32.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0
ISO 631
-9.50
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 18.0
72
34.00 -11.20
-11.60

33.00
-12.00
32.00
-12.40
31.00
R2 = 0.0124
-12.80
30.00
-13.20
29.00 30.00 30.50 31.00 31.50 32.00 32.50 33.00
0.0 5.0 10.0 15.0 20.0
ISO 620
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
0.0 5.0 10.0 15.0 20.0
34.00 -11.80
33.50 -12.00
33.00
-12.20
32.50
-12.40
32.00
R2 = 0.0003
31.50 -12.60
31.00 -12.80
30.50
-13.00
30.00 30.00 30.50 31.00 31.50 32.00 32.50 33.00 33.50 34.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
ISO 619
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
73
32.00 -10.60
31.50 -10.80

δ18O bioapatite (‰) 31.00 -11.00
30.50 -11.20
30.00 -11.40
29.50 R2 = 0.752
-11.60
29.00 -11.80
28.50 -12.00
28.00 28.00 28.50 29.00 29.50 30.00 30.50 31.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 δ18O bioapatite (‰)
ISO 628
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0
32.00 -12.20
-12.40
31.50
-12.60
31.00
-12.80
R2 = 0.172
30.50 -13.00
30.00 -13.20
-13.40
29.50 30.60 30.70 30.80 30.90 31.00 31.10 31.20 31.30 31.40
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
ISO 609
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
-14.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
74
32.50 -11.40
32.00 -11.80

-12.20
31.50
-12.60
31.00 R2 = 0.6406
-13.00
30.50 -13.40
-13.80
30.00
-14.20
29.50 30.00 30.50 31.00 31.50 32.00 32.50
0.0 5.0 10.0 15.0 20.0
ISO 608
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
-14.50
0.0 5.0 10.0 15.0 20.0
31.50 -10.80
-11.00
31.00
-11.20
30.50 -11.40
-11.60
30.00 R2 = 0.6798
-11.80
29.50 -12.00
-12.20
29.00
-12.40
28.50 28.50 29.00 29.50 30.00 30.50 31.00 31.50
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0 δ18O bioapatite (‰)
ISO 613
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
75
31.00 -11.20
30.50 -11.40

-11.60
30.00
-11.80
29.50
-12.00
29.00 R2 = 0.1978
-12.20
28.50
-12.40
28.00 28.00 28.50 29.00 29.50 30.00 30.50 31.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 δ18O bioapatite (‰)
ISO 616
-10.00
-10.50
-11.00
-11.50
-12.00
-12.50
-13.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
31.50 -10.00
31.00
-10.20
30.50 -10.40
30.00
-10.60
29.50 R2 = 0.051
-10.80
29.00
-11.00
28.50 28.50 29.00 29.50 30.00 30.50 31.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 δ18O bioapatite (‰)
ISO 615
-9.00
-9.50
-10.00
-10.50
-11.00
-11.50
-12.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0
76
29.00 -12.00
28.50

-12.40
28.00 -12.80
27.50
-13.20
27.00
-13.60
26.50 R2 = 0.6271
-14.00
26.00 27.20 27.40 27.60 27.80 28.00 28.20 28.40 28.60
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
ISO 605
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0
31.50 -12.50
-12.70
31.00
30.50 -12.90
R2 = 0.0828
-13.10
30.00
-13.30
29.50
-13.50
29.00 29.60 29.80 30.00 30.20 30.40 30.60 30.80 31.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
ISO 604
-12.00
-12.40
-12.80
-13.20
-13.60
-14.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 14.0 16.0
77
35.00 -12.20
-12.40

34.50
-12.60
34.00
-12.80
33.50 R2 = 0.1768
-13.00
33.00
-13.20
32.50
-13.40
32.00 33.40 33.50 33.60 33.70 33.80
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 δ18O bioapatite (‰)
ISO 602
-11.00
-11.50
-12.00
-12.50
-13.00
-13.50
-14.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0
31.50 -12.60
31.00 -12.80
-13.00
30.50 R2 = 0.0225
-13.20
30.00 -13.40
29.50 -13.60
-13.80
29.00
-14.00
28.50 29.00 29.50 30.00 30.50 31.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 δ18O bioapatite (‰)
ISO 692
-12.00
-12.50
-13.00
-13.50
-14.00
-14.50
-15.00
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0
78
33.00 -12.20
-12.40

32.00 -12.60
-12.80
R2 = 0.7797
31.00 -13.00
-13.20
30.00 -13.40
-13.60
29.00 -13.80
-14.00
28.00 28.00 29.00 30.00 31.00 32.00 33.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0 δ18O bioapatite (‰)
ISO 689
-12.00
-12.40
-12.80
-13.20
-13.60
-14.00
0.0 2.0 4.0 6.0 8.0 10.0 12.0
32.00 -11.60
31.90 -11.80
-12.00
31.80
-12.20
31.70 R2 = 0.7852
-12.40
31.60
-12.60
31.50
-12.80
31.40 -13.00
31.30 31.30 31.40 31.50 31.60 31.70 31.80 31.90 32.00
0.0 1.0 2.0 3.0 4.0 5.0 δ18O bioapatite (‰)
ISO 601
-11.80
-11.90
-12.00
-12.10
-12.20
-12.30
-12.40
-12.50
-12.60
0.0 1.0 2.0 3.0 4.0 5.0
79
HOL 689 Covariance HOL 689
32.00 -8.00
31.00 -8.50

30.00 -9.00
R2 = 0.0015
-9.50
29.00
-10.00
28.00
-10.50
27.00
-11.00
26.00 27.00 28.00 29.00 30.00 31.00
0.0 10.0 20.0 30.0 40.0 50.0 60.0 δ18O bioapatite (‰)
HOL 689
-6.00
-7.00
-8.00
-9.00
-10.00
-11.00
-12.00
0.0 10.0 20.0 30.0 40.0 50.0 60.0
34.00 -7.00
33.00 -7.50
32.00 -8.00
31.00 -8.50 R2 = 0.6018
30.00
-9.00
29.00
-9.50
28.00
-10.00
27.00 28.00 29.00 30.00 31.00 32.00 33.00
0.0 5.0 10.0 15.0 20.0 25.0 δ18O bioapatite (‰)
HOL 753
-6.00
-7.00
-8.00
-9.00
-10.00
-11.00
0.0 5.0 10.0 15.0 20.0 25.0
80
33.00 -8.00
32.50

-8.50
32.00 R2 = 0.0787
-9.00
31.50
31.00 -9.50
30.50 -10.00
30.00
-10.50
29.50 29.50 30.00 30.50 31.00 31.50 32.00 32.50 33.00
0.0 5.0 10.0 15.0 20.0 25.0 30.0 δ18O bioapatite (‰)
HOL 370
-8.00
-8.50
-9.00
-9.50
-10.00
-10.50
-11.00
0.0 5.0 10.0 15.0 20.0 25.0 30.0
32.00 -7.50
31.00 -8.00
30.00 -8.50
29.00 -9.00
28.00 -9.50
R2 = 4E-05
-10.00
27.00
-10.50
26.00
26.00 27.00 28.00 29.00 30.00 31.00 32.00
0.0 10.0 20.0 30.0 40.0 50.0
HOL 154
-7.00
-8.00
-9.00
-10.00
-11.00
0.0 10.0 20.0 30.0 40.0 50.0
81
Appendix 3
Modern Plant Sample Data, Israel
Common
Sample Region Family Genus Species name δ13C (‰)
IS-1 Lod Junction Anacardiaceae Pistacia lentiscus Mastic tree -27.7
IS-2 Lod Junction Leguminosae Ceratonia siliqua Carob -27.1
IS-3 Lod Junction Pinaceae Pinus halepensis pine -26.5
IS-4 Lod Junction Rhamnaceae Rhamnus palaestinus buckthorn -26.8
IS-5 Lod Junction Cupressaceae Juniperus phoenica Juniper -25.3
IS-6 Lod Junction Moraceae Ficus carica Fig -25.5
IS-7 Lod Junction Anacardiaceae Pistacia saportae pistachio? -28.6
IS-8 Lod Junction Leguminosae Ceratonia siliqua Carob -28.5
IS-9 Lod Junction Cupressaceae Cupressus sempervirens Italian Cypress -29.4
IS-10 Lod Junction Santalaceae Osyrus alba -27.7
IS-11 Lod Junction Rosaceae Sarcopoterium spinosum shrub -25.1
IS-12 Lod Junction Leguminosae Acacia tortilis? acacia -26.9
IS-13 Lod Junction Capparidaceae Capparis ovata caper -29.9
IS-14 Lod Junction Liliaceae Asparagus aphyllus Lily -28.5
IS-15 Lod Junction Gramineae Triticum? grass -28.9
IS-16 Lod Junction Gramineae Poa/Puccinellia grass -28.5
IS-17 Lod Junction Gramineae Panicum turgidum? grass -30.8
IS-18 Lod Junction Gramineae Dactylis grass -25.3
IS-19 Lod Junction Gramineae Bromus grass -27.6
IS-20 Lod Junction Santalaceae Osyrus alba -29.4
IS-21 Lod Junction Labiatae Marrubium/Origanum? herb -28.6
IS-22 Lod Junction Unknown -27.4
IS-23 Lod Junction Compositae -27.0
IS-26 Lod Junction Compositae Carlina hispanica? Thistle -27.5
IS-27 Lod Junction Compositae Carlina racemosa? Thistle -27.2
IS-28 Lod Junction Compositae Picnomon? -24.4
IS-29 Lod Junction Compositae Carlina curetum? Thistle -25.8
Spanish
IS-30 Lod Junction Leguminosae Spartium junceum? broom -30.0
IS-31 Lod Junction Umbellaferae Pimpinella/Pituranthos? aniseseed -26.7
IS-32 Lod Junction Unknown -29.4
IS-33 Lod Junction Umbellaferae Daucus carota? wild carrot -24.5
IS-34 Lod Junction Caryophyllaceae Dianthus cyri? carnation -27.7
IS-35 Mt Carmel Anacardiaceae Pistacia lentiscus Mastic tree -27.1
IS-36 Mt Carmel Anacardiaceae Pistacia lentiscus Mastic tree -27.1
IS-37 Mt Carmel Rosaceae Cerasus microcarpa cherry -28.2
IS-38 Mt Carmel Unknown -24.6
IS-39 Mt Carmel Leguminosae Ceratonia siliqua Carob -32.2
ithaburensis?
IS-40 Mt Carmel Fagaceae Quercus (type 2) oak -28.2
IS-41 Mt Carmel Fagaceae Quercus boissieri oak -28.0
IS-42 Mt Carmel Liliaceae Smilax aspera sasparilla -24.0
Sarcpoterium
IS-43 Mt Carmel Rosaceae (Poterium) spinosum shrub? -24.0
IS-44 Mt Carmel Unknown Hypericum hyssopifolium St. John's wort -28.4
82
IS-45 Mt Carmel Leguminosae Calycotome villosa herb? -24.5
IS-46 Mt Carmel Liliaceae Asparagus acutifolius Lily -25.6
IS-47 Mt Carmel Unknown Rapistrum rugosum herb -28.2
IS-49 Mt Carmel Gramineae Bromus grass -24.7
IS-50 Mt Carmel Gramineae Stipa? grass -29.5
IS-52 Mt Carmel Malvaceae Abutilon theophrasti China jute -25.8
IS-53 Mt Carmel Unknown Rumex pulcher? sorrel -27.7
IS-56 Mt Carmel Compositae Carlina hispanica? Thistle -27.3
IS-57 Mt Carmel Compositae Carlina curetum? Thistle -26.7
83

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INTER- AND INTRA-TOOTH ISOTOPIC VARIATION IN

MAMMALIAN TOOTH ENAMEL FROM WESTERN ISRAEL:

A Prepublication Manuscript Submitted to the Faculty of the

In Partial Fulfillment of the Requirements

In the Graduate College

APPROVAL BY RESEARCH COMMITTEE

As members of the Research Committee, we recommend that this prepublication

Neolithic tooth assemblages (Shahack-Gross et al., 1999; Richards et al., 2003).

1998; Zazzo et al., 2002; Balasse et al., 2003).

Considerable turnover has been noted in mammalian communities of the region

could be potentially explained by 1) changes in human subsistence behavior and prey

selection, or by 2) climate-driven environmental changes. Here we use stable carbon and

of the human evolutionary record in the region, it is critical to closely examine

movements of early modern human populations into the Levant.

Site Locations, Chronologies, and Regional Climate

archaeological and natural deposits, including extensive calcite flowstones. Uranium

series (230Th/234U/238U) dating of a massive flowstone and a speleothem crust that

most of the Acheulo-Yabrudian layers. The flowstone is directly overlain by a thin

archaeological deposit, which accumulated during a hiatus in speleothem deposition from

The ~207 ka BP age estimate for the upper Acheulo-Yabrudian (Layer 2) is

stratigraphy (Layers 3 and 4) is best described as provisional. Barkai et al. (2003)

groups of 300-350 ka BP and 200-215 ka BP.

of Israel, approximately 30 km northeast of Haifa, at an elevation of ~250 m asl. The two

terminal Upper Paleolithic and early Epipaleolithic periods. Middle Paleolithic

chronology at Hayonim has been determined by uranium-series dating of speleothems

All chronologies are presented in Table 1.

interspersed with stands of wild wheat and barley (Zohary, 1973).

Tooth Enamel Formation and Stable Isotope Analysis

Tooth enamel (bioapatite) is highly resistant to isotopic alteration compared to

Gross et al., 1999). It is composed of tightly packed crystallites of biogenic

hydroxyapatite (Ca10[PO4,CO3]6[OH]2) that are ~50-100 nm in diameter and >1000nm in

along the growth axis of a mammalian tooth, representing a partially time-averaged

Balasse et al., 2003).

body water is represented by:

water (δ18Olw) can be evaporatively enriched in 18O by 10-25‰ in comparison to stem

and local meteoric water (Dongmann et al., 1974).

The carbon isotope composition of mammalian tooth enamel (δ13Cbioapatite) is

through time in δ13Cbioapatite values is to be expected, although minor variations may

1999; Passey et al., 2005).

Fallow deer (Dama mesopotamica) and mountain gazelle (Gazella gazella)

issue will be discussed in more detail below.

carbon (δ13C) isotope measurements of the structural carbonate component of bioapatite

binocular microscope with a 0.9 mm diamond-tipped drill bit. Furrows approximately 1

ensured that dentine was excluded from the sample.

Powdered enamel is typically pretreated with dilute sodium hypochlorite (NaOCl)

a) no pretreatment; b) 2% NaOCl for 12 hours; c) 2% NaOCl for 12 hours, followed by

samples of each aliquot were analyzed.

The δ13Cbioapatite and δ18Obioapatite values were measured using an automated

carbonate preparation device (KIEL-III) coupled to a gas-ratio mass spectrometer

an automated CHN analyzer (Costech) coupled to a continuous-flow mass spectrometer

matter are reported relative to VPDB.

Previous pretreatment experiments on tooth enamel (Lee-Thorpe & van der

procedures have on enamel of various ages, and to determine if pretreatment is necessary

mentioned above is most relevant to this discussion (Figs. 2a and 2b).

enamel pretreatment procedure is unnecessary in the context of our investigation (as in

Zazzo et al., 2005).

Temporal trends in isotopic values

20 ka BP, after which time they decrease.

The isotopic values of intra-tooth enamel microsamples vary substantially. The

4.1‰, respectively. Overall, intra-tooth isotopic variation in δ18Obioapatite values is greater

“seasonal” range of δ18Obioapatite values (4.1‰) is similar to the seasonal variation in

of δ18Obioapatite values (2.3‰) is about half that.

Local Vegetation and Meteoric Water

The wild vegetation surrounding the three archaeological sites is composed