Quaternary Research 63 (2005) 15 – 30

www.elsevier.com/locate/yqres

Environmental significance of 13C/12C and 18O/16O ratios of modern

land-snail shells from the southern great plains of North America
Meena Balakrishnana, Crayton J. Yappa,*, James L. Thelerb,
Brian J. Carterc, Don G. Wyckoff d
a
Department of Geological Sciences, Southern Methodist University, Dallas, TX 75275-0395, USA
b
Department of Sociology and Archaeology, University of Wisconsin-La Crosse, La Crosse, WI 54601, USA
c
Department of Plant and Soil Sciences, Oklahoma State University, Stillwater, OK 74078, USA
d
Oklahoma Museum of Natural History, University of Oklahoma, Norman, OK 73019, USA

Received 11 March 2004

Available online 26 November 2004

Abstract
13 12
C/ C and 18O/16O ratios of aragonite shells of modern land snails from the southern Great Plains of North America were measured for
samples from twelve localities in a narrow east–west corridor that extended from the Flint Hills in North Central Oklahoma to the foothills of
the Sangre de Cristo Mountains in Northern New Mexico, USA. Across the study area, shell y18O values (PDB scale) ranged from
4.1x to
1.2x, while y13C values ranged from
13.2x to 0.0x. y18O values of the shell aragonite were predicted with a published, steady state,
evaporative flux balance model. The predicted values differed (with one exception) by less than 1x from locality averages of measured y18O
values. This similarity suggests that relative humidity at the time of snail activity is an important control on the y18O values of the aragonite
and emphasizes the seasonal nature of the climatic information preserved in the shells. Correlated y13C values of coexisting Vallonia and
Gastrocopta suggest similar feeding habits and imply that these genera can provide information on variations in southern Great Plains plant
ecology. Although there is considerable scatter, multispecies, transect average y13C values of the modern aragonite shells are related to
variations in the type of photosynthesis (i.e., C3, C4) in the local plant communities. The results of this study emphasize the desirability of
obtaining isotope ratios representing averages of many shells in a locale to reduce possible biases associated with local variations among
individuals, species, etc., and thus better represent the bneighborhoodQ scale temporal and/or spatial environmental variations of interest in
studies of modern and ancient systems.
D 2004 University of Washington. All rights reserved.

Keywords: Land snails; Oxygen isotopes; Carbon isotopes; Climate; Relative humidity; C3 plants; C4 plants

Introduction y18O of ambient meteoric water (e.g., Balakrishnan and

Studies of the isotopic compositions of the aragonitic
shells of land snails from different parts of the world attest
to their value as environmental indicators (Goodfriend and
Ellis, 2002; Goodfriend and Magaritz, 1987; Goodfriend et
al., 1989; Lécolle, 1985; Magaritz and Heller, 1980, 1983;
Magaritz et al., 1981; Yapp, 1979). y18O values of these
shells have been related to local climatic parameters and the
* Corresponding author. Fax: +1 214 768 2701.
E-mail address: cjyapp@mail.smu.edu (C.J. Yapp).
0033-5894/$ - see front matter D 2004 University of Washington. All rights reserved.
doi:10.1016/j.yqres.2004.09.009
Yapp, 2004; Goodfriend et al., 1989; Lécolle, 1985;
Magaritz and Heller, 1980; Magaritz et al., 1981; Yapp,
1979). y13C values of shell aragonite may be correlated with
the y13C of local vegetation (e.g., Balakrishnan and Yapp,
2004; Goodfriend and Magaritz, 1987; Metref et al., 2003;
Stott, 2002).
Goodfriend and Ellis (2002) conducted a study of the
y13C and y18O variations of shell aragonite and shell organic
matrix in two species of land snails of the genus Rabdotus
along a transect from Northeast to Southwest Texas, USA.
They examined evidence for relationships between ambient
environmental parameters and the y13C or y18O values of
16 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30
shell aragonite. y13C values of shell aragonite were
correlated with y13C values of organic matrix in the shell,
which suggests that the shell y13C reflects diet with an offset
associated with the snail physiology and equilibrium and
kinetic fractionation processes. Relationships of y18O
variations of Rabdotus to the environmental variables
discussed by Goodfriend and Ellis (2002) appeared to be
more problematic than for carbon.
In the current paper, we present measurements of
variations in the carbon and oxygen isotope compositions
of shells of other species of modern land snails from a
different portion of the southern Great Plains of North
America. Samples in this study are described in the work of
Theler et al. (2004) and are from a narrow east–west
corridor extending across much of Oklahoma and into
Northeastern New Mexico, USA. The data are discussed in
terms of their relationship to modern environmental
variables to examine the likelihood that isotopic data from
ancient land snails in the region might have paleoenvir-
onmental significance.
Samples and study area
Samples
The modern snail population procured for this study
formed part of an extensive terrestrial gastropod survey
(Theler et al., 2004; Wyckoff et al., 1997) in the southern
Great Plains of North America. Samples were collected at
12 localities along an east–west corridor extending from
North Central Oklahoma to Northeastern New Mexico
between 36826VN to 36858VN latitude and 96849VW to
104857VW longitude (Theler et al., 2004; Wyckoff et al.,
1997). The site names and locations are depicted in Figure
1. The study corridor is 640 km long east to west and about
100 km wide and extends from the Flint Hills of North
Central Oklahoma to the foothills of the Sangre de Cristo
Mountains, near Cimarron, New Mexico (Fig. 1). Elevation
in the study area slowly increases from 330 m above sea
level at the eastern end of the corridor to about 2290 m
above sea level at the western end.
Vegetation
There are four distinct biotic districts in the sampled area
(Fig. 1). Mixed-grass plains occupy the eastern section
(Blair and Hubbell, 1938; Carpenter, 1940; Ostlie et al.,
1997; Shelford, 1963). This district marks the transition
from tall grass prairie in the east to short grass prairie in the
west and is characterized by several species of Ascoparius
(Ascoparius saccharoides, Ascoparius furcatus, Ascoparius
smithii), grama grasses (Bouteloua gracilis, Bouteloua
racemosa, Bouteloua hirsuta, and Bouteloua curtipendula),
and buffalo grass (Buchloë dactyloides) (Blair and Hubbell,
1938; Bruner, 1931; Carpenter, 1940; Kuchler, 1964; Ostlie
et al., 1997; Risser, 1985, 1990; Shelford, 1963; Weaver and
Albertson, 1956).
The second biotic region, short grass prairie (immedi-
ately to the west of the mixed-grass prairie, Fig. 1), contains
vegetation that includes buffalo grass, hairy grama (B.
hirsuta), blue grama (B. gracilis), and western wheatgrass
(Pascopyrum smithii) (Blair and Hubbell, 1938; Bruner,
1931; Carpenter, 1940; Kuchler, 1964; Ostlie et al., 1997;
Risser, 1990; Shelford, 1963).
A third biotic district, the pinyon–juniper shrub grass-
land, Raton subsection (Blair and Hubbell, 1938) extends
from the northwest corner of the panhandle into the
northeastern corner of New Mexico (Fig. 1). Vegetation in
this region includes junipers (Juniperus monosperma,
Juniperus osteosperma), oak (Quercus mohriana), and pine
(Pinus edulis, Pinus monophylla), along with plants such as
silverbeard grass (A. saccharoides), blue grama, and some
species of prickly pear cactus (Opuntia sp.) (Blair and
Hubbell, 1938; Ostlie et al., 1997).
The dry pine forest at the western end of the study area
(Fig. 1) is not strictly a Plains ecosystem (Shelford, 1963).
Vegetation includes oak and pine along with hairy and blue
grama grasses (Shelford, 1963; Theler et al., 2004; Wyckoff
et al., 1997).
Climate and isotopes in precipitation
Average annual precipitation on the southern plains
generally decreases from over 1020 mm/yr in the east to
255 mm/yr in the west (Ostlie et al., 1997). The mean
annual temperatures in the mixed-grass prairie range from
158 to 178C, while the corresponding mean annual
precipitation ranges from 670 to 790 mm (Blair and
Hubbell, 1938). Annual temperatures in the short grass
prairie range from 128 to 138C and mean annual precip-
itation from 450 to 560 mm (Blair and Hubbell, 1938).
Average annual temperatures in the dry pine forest of the
foothills of the Sangre de Cristo mountains are about 118C
and average rainfall is about 400 mm (Climate Data Center,
New Mexico State University at www.weather.nmsu.edu).
Precipitation in these regions derives principally from three
locations (Elliot, 1949; Nativ and Riggio, 1990).
From late March into July, moisture is derived primarily
from the Gulf of Mexico (curve ba,Q Fig. 2). From October
to early March, the Northern Pacific Ocean is the primary
moisture source (Nativ and Riggio, 1990) (curve bc,Q Fig.
2). In the western end of the study area (Northeastern New
Mexico), both the Gulf of Mexico and the central Pacific
Ocean contribute moisture during the middle to late summer
and early fall (curves baQ and bb,Q Fig. 2), although
contributions from the Gulf of Mexico are predominant.
The central Pacific component of this moisture is brought in
by the Mexican monsoons (Douglas et al., 1993; Nativ and
Riggio, 1990).
The temperature and relative humidity at which ocean
water evaporates, the air mass history, and the local
 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30 17

Figure 1. Map of the study area in the southern Great Plains of North America indicating the collection localities and the major ecological regions. Localities: 1
Kubic; 2 Bluff Creek; 3 Salt Fork; 4 McDaniel; 5 Burnham; 6 Big Salt Plain; 7 Skull Springs; 8 Hitch; 9 Black Mesa; 10 Owensby; 11 C. S. Ranch; 12 Chase
(data sources: Blair and Hubbell, 1938; Carpenter, 1940; Shelford, 1963; Wyckoff et al., 1997; Theler et al., 2004).

temperature of condensation are among the controls on Experimental

the isotopic composition of precipitation (Dansgaard,
1964; Rozanski et al., 1993). The different air masses
that contribute precipitation to the southern Plains impart
seasonally distinct y18O values to the rain and/or snow.
y18O values of summer rains originating from Gulf of
Mexico moisture are more positive than those of winter
precipitation originating from the North Pacific Ocean
(Nativ and Riggio, 1990). This is primarily a consequence
of the higher elevations (lower temperatures) and longer
transport distances experienced by air masses from the
North Pacific (Nativ and Riggio, 1990; Rozanski et al.,
1993).
Sampling systematics
Results of an initial survey, collection, and documentation
of the modern gastropods in this region are found in Wyckoff
et al. (1997) and Theler et al. (2004). These include
description of the dominant vegetation types at each locality.
Collections of the snails analyzed for the current work were
made at 12 localities along the 640-km sample corridor at
sites for which living snails could be expected (for sample
collection rationale, see Theler et al., 2004). Each sample
locality was restricted to a circle with a diameter of ~400 m
18 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30

Figure 2. Source regions and generalized trajectories of major moisture-bearing air masses that bring precipitation to the southern Great Plains of North
America. Trajectory: (a) March into July or August; (b) middle to late summer and early fall; (c) October to early March (see text). mP = maritime polar air
mass and mT = maritime tropical air mass (after Elliot, 1949; Nativ and Riggio, 1990). The shaded rectangle encompasses the land snail sites of Figure 1.

(Theler et al., 2004; Wyckoff et al., 1997). Each circle
contained several niches. Niches (recognized by differences
in vegetation, slope, and aspect) favorable for snail habitation
were sampled along a linear btransectQ (Theler et al., 2004;
Wyckoff et al., 1997) that was usually not more than 100 m in
length. Along each transect, there were three sample
collection sites (usually 10–20 m apart) called breplicationsQ
A, B, and C (Wyckoff et al., 1997). The term bsampleQ was
substituted for breplicationQ by Theler et al. (2004). However,
in the current work, we will retain breplicationQ (sensu;
Wyckoff et al., 1997) to avoid confusion with our more
generic use of the term bsample,Q which refers herein to any
collected material of interest. Thus, as an example, the
sample locality bHitchQ has three transects (Hitch 21, Hitch
22, and Hitch 23) with each transect in Hitch in turn
comprised of three replications (e.g., Hitch 21A, Hitch 21B,
and Hitch 21C). There is a total of 38 transects among the 12
localities, and a total of 114 breplicationsQ summed over all of
the 38 transects (i.e., 3  38). At each breplication,Q lower
parts of growing vegetation, decaying vegetation, and 2 cm
of topsoil from a 50  50 cm area were collected (Wyckoff et
al., 1997; Theler et al., 2004). Wyckoff et al. (1997) and
Theler et al. (2004) sieved and sorted these samples in the
laboratory to extract the snail shells, followed by identifica-
tion and population analyses.
Because of the destructive nature of the isotopic analyses
and because it was necessary to retain snail shells for
archival purposes, only those from sample sites represented
by large numbers of collected shells were analyzed. Hence,
of the total of 114 breplicationsQ among the 38 transects,
only 71 replications from 34 transects are represented in this
study. Most of the samples were collected in 1995
(collection from the Owensby site was made in 1996) and
were alive at the time or within 1 yr of collection (Theler et
al., 2004; Wyckoff et al., 1997).
Selection of species for isotopic analyses
Vallonia and Gastrocopta were the principal genera
employed for isotopic analyses of snail shells because they
are present throughout most, or all, of the study area (Theler
et al., 2004; Wyckoff et al., 1997). For some localities,
snails representing a few other genera were also isotopically
analyzed, but these other genera did not have the wide
distribution exhibited by Vallonia or Gastrocopta. The
genus Vallonia is represented by the species Vallonia
parvula at the lower elevations and Vallonia gracilicosta
at the higher elevations. The latter is often associated with
deposits of Pleistocene age in the southern Great Plains
(Rossignol et al., 2004; Theler et al., 2004; Wyckoff et al.,
1997). Vallonia juveniles were also analyzed but could not
be identified at the species level (Theler et al., 2004;
Wyckoff et al., 1997). The genus Gastrocopta was
represented by Gastrocopta contracta, Gastrocopta holzin-
geri, Gastrocopta pentodon, Gastrocopta armifera, Gastro-
copta cristata, Gastrocopta pilsbryana, Gastrocopta
 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30 19

procera, and Gastrocopta pellucida. The two latter species

exhibit relatively large distribution ranges. However, G.
procera was notably absent at the higher altitudes in the
survey (Theler et al., 2004; Wyckoff et al., 1997).

Laboratory preparations and analyses

Each isotopically analyzed sample consisted of multiple

entire shells of adults of a particular species (or genus for
juveniles of Vallonia) from a breplication.Q The shells were
initially rinsed with deionized water, then treated ultrasoni-
cally in deionized water to remove any adhering particles of
organic matter or other debris. The shells were gently crushed
and treated with 5% reagent-grade sodium hypochlorite at
room temperature for about 7–8 h to remove organic matter.
They were then rinsed thoroughly with deionized water and
dried in air at about 40–508C. The shell fragments were
subsequently reacted overnight in vacuum with 100% H3PO4
at 258C following the method of McCrea (1950). The CO2
prepared from the shell aragonite was analyzed for y13C and
y18O on a Finnigan MAT 252 mass spectrometer. Analytical
uncertainty is about F0.1x. The y values are defined as:


y13 C or y18 O ¼ R sample =R standard
1  1000 x

where R = 13C/12C or 18O/16O. y13C and y18O are reported

relative to the PDB standard (Craig, 1957).

Results and discussions
Ranges of d18O and d13C of aragonite in land-snail shells
Figure 3. Ranges of Southern Great Plains land snails and snail shell y18O
and y13C values measured for breplicationsQ in the current study compared
with various published ranges (see text). Open portion of the range of North
American y18O values encompasses the values reported for shells from a
carnivorous, cold-tolerant snail (see text for reference).

y18O and y13C values were measured for 162 samples
from the 12 localities of Figure 1 (see Appendix A). y18O
values of the aragonite shells of these modern snails ranged
from
4.1x to 1.2x (Fig. 3). Published y18O values of
shells from globally distributed modern land snails that are
wholly subaerial (as opposed to semiaquatic) range from

11.7x to 4.5x (Goodfriend and Ellis, 2002; Goodfriend
and Magaritz, 1987; Goodfriend et al., 1989; Lécolle, 1985;
Magaritz and Heller, 1980; Magaritz et al., 1981; Sharpe et
al., 1994; Yapp, 1979). In North America, reported y18O
values of land-snail shells range from
11.7x to 0.2x
(Goodfriend and Ellis, 2002; Sharpe et al., 1994; Yapp,
1979). If the results for a carnivorous, cold-tolerant snail of
the genus Vitrina (Sharpe et al., 1994) from Deer Creek,
Nevada, are excluded, y18O values analyzed to date for
North America range from
5.8x to 0.2x. The current
work extends the upper range of North American values by
1.0x (Fig. 3).
Previous studies of y13C values of shell aragonite from
snails in their natural settings reported values ranging from

13.5x to 0.5x (Goodfriend and Ellis, 2002; Goodfriend
and Magaritz, 1987; Lécolle, 1983, 1984; Magaritz and
Heller, 1980, 1983; Magaritz et al., 1981; Yapp, 1979). For
North America, published y13C values range from
12.5x
to
2.5x (Goodfriend and Ellis, 2002; Yapp, 1979). The
y13C values of snail shell aragonite studied for the current
work, range from
13.2x to 0.0x (Fig. 3). By contrast,
y13C values of shell aragonite of experimentally cultured
Helix aspersa that were fed a controlled diet ranged from

24.3x to 2.5x (Stott, 2002). Controlled experiments also
indicate differences in y13C values among adults, hatched,
and 1-month-old individuals of H. aspersa fed diets with
identical y13C values (Metref et al., 2003).
Snail shell isotopic variation among populations
Variations in the isotopic composition of snail shells exist
among replications within a transect and also among species
in a replication (Appendix A). Within a sample locality,
small-scale variations in topography, vegetation, local
moisture availability, snail ages, physiology, etc., are
expected. Such variations might contribute to the observed
scatter in isotopic ratios among genera, species, or
individual samples.
Isotopic analyses of Vallonia and Gastrocopta total 143
and represent the majority of the analyzed shell samples, but
20 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30
Figure 4. Southern Great Plains land snails. Comparison of average
isotopic compositions of the aragonitic shells of Gastrocopta and Vallonia
from the same breplicationsQ (see text). (a) comparison of y18O values and
(b) comparison of y13C values. Dashed line in panel a is the reference line
y = x. Solid line in panel b is the linear regression of the data with the
corresponding equation, r 2 and P.
there were also analyses (numbers in parentheses) of
bunderrepresentedQ genera as follows (see Appendix A):
Pupoides (9), Succineidae (3), Pupilla (2), Cionella (1),
Zonitoides (1), Helicodiscus (1), Glyphyalinia (1), and
Hawaiia (1). The ranges of y18O values for Vallonia and
Gastrocopta are about 2.8x and 2.0x, respectively (Fig.
4a). Corresponding y18O values of Vallonia and Gastrocopta
Figure 5. Southern Great Plains land snails. Comparison of y18O of the aragonitic shells of other species used in this study with that of coexisting (a) Vallonia
and (b) Gastrocopta in the breplicationsQ. (c) Comparison of y13C values of the shell aragonite of other species with those of coexisting Vallonia and/or
Gastrocopta in the same breplicationsQ (see text). Dashed lines are the reference lines y = x.
indicate no correlation among the various sites (Fig. 4a). A
similar lack of correlation was evident when y18O values of
underrepresented genera were compared with values for
coexisting Vallonia (Fig. 5a) or Gastrocopta (Fig. 5b). The
particular origins of these differences are unknown. Without
studies under controlled age and environmental conditions
(e.g., Metref et al., 2003; Stott, 2002), distinguishing micro-
environmental effects from the effects of age and/or species-
related physiological differences is not possible at present.
The respective ranges of y13C values for Vallonia and
Gastrocopta shell aragonite are each about 8–9x, and
samples from the same breplicationsQ show a significant
correlation (Fig. 4b). This correlation could indicate similar
feeding habits in both genera, making them potential sources
of isotopic information on the paleoecology of a locale.
Because of this correlation of Vallonia and Gastrocopta y13C
values, y13C values of eight other genera that coexisted with
one or both of these genera in the various breplicationsQ are
plotted against y13C of either Vallonia or Gastrocopta in a
single figure (Fig. 5c). There are insufficient data to establish
whether the y13C values of any of the bunderrepresentedQ
genera might be correlated with Vallonia or Gastrocopta, but
overall there is considerable scatter among the species.
Additional comparative analyses of, and controlled experi-
ments on, these genera are needed.
An exploratory evaluation of the effect of life histories
and ages on the isotopic composition of snails was
attempted using the data for adult and juvenile Vallonia
(see Appendix A). y18O values of shells of coexisting adult
and juvenile Vallonia from the breplicationsQ are not
correlated (Fig. 6a). The y18O values of the adult snails
represent averages over longer periods of time than those of
the juveniles. Short time scale environmental conditions that
affect the y18O values of the body fluid in juveniles should
determine the y18O values of the shell aragonite and these
conditions might contrast with the longer term-averages of
the adults.
The absence of a correlation in the y18O data of Figure 6a
contrasts with the good correlation of the y13C values of the
adults and juveniles in Figure 6b. The y13C correlation
could indicate that, in these locales, the carbon isotope
 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30
Figure 6. Comparison of coexisting adults and juveniles of southern Great
Plains Vallonia in breplicationsQ (see text). (a) y18O values of aragonite
shells and (b) y13C values. Solid line in panel b is the linear regression of
the data with the corresponding equation, r 2 and P.
composition of vegetation (as well as snail feeding patterns)
is not as variable on small scales as the oxygen isotope
composition of ambient moisture. However, the apparent
contrast in the respective oxygen and carbon isotope
relationships (adults or juveniles, Figs. 4 and 6) might also
be, in part, a consequence of the smaller ranges for the y18O
values. Thus, if the magnitude of the normal scatter in the
breplicationsQ data were comparable to the smaller total
range of the y18O values, it would mask any evidence for a
broader y18O correlation among genera or age groups.
Snail shell isotope compositions and environmental
parameters
Carbon isotopes
As noted, vegetation in the study area is dominantly
grasslands at lower elevations and trees at higher
elevations (Fig. 1). Based on the nature of the photo-
synthetic pathway, plants are classified as C3, C4, or CAM
(e.g., Jacobs et al., 1999). y13C values of C3 plants range
from
33x to
21 x, while y13C values of C4 plants
typically range from
17x to
9x (e.g., Cerling and
21
Quade, 1993). CAM plants have y13C values that range
between the values for C3 and C4 plants (e.g., Cerling and
Quade, 1993). Most of the grasses in the southern Great
Plains are C4 plants (Tieszen et al., 1997), and the trees
and most of the forbs are C3 plants (Watson and Dallwitz,
http://biodiversity.uno.edu/delta/).
If snail shell y13C values are influenced by the y13C of
their diets (e.g., Balakrishnan and Yapp, 2004; Francey,
1983; Goodfriend and Ellis, 2002; Goodfriend and Magar-
itz, 1987; Metref et al., 2003; Stott, 2002), the spatial
transitions in the ecology of the study corridor suggest that
the y13C values of the shells should be more negative at the
higher elevations toward the west. y13C values of the shell
aragonite from all breplicationsQ (Appendix A) and their
averages for each transect (Table 1) are plotted against
elevation in Figures 7a and 7b, respectively. There is
considerable scatter, but some suggestion of the expected
general shift to lower y13C values as altitude increases (Fig.
7). However, the regional vegetation trend does not explain
the very large range of snail shell y13C values at the lowest
elevations on the eastern end of the sample corridor (Figs.
7a and 7b). Such a large range may be explained, in part, by
local variations in proportions of C3 and C4 plants.
Wyckoff et al. (1997) and Theler et al. (2004)
identified plant species at each sample site in the study
corridor. For the current work, their identifications (no
quantitative estimates of type) were used to classify the
vegetation in each transect as C3-dominant, C4-dominant,
or mixed, including CAM (Ehleringer et al., 1997;
Owensby et al., 1997; Sage et al., 1999; Appalachian
Farming Systems Research Center, http://www.arserrc.gov/
beckley/C3C4LIST.htm). The classifications are presented
in Table 1.
The average y13C values of the land-snail shells from
each transect are listed in Table 1 and plotted in Figure 8
against the corresponding classification of local vegetation,
which is arranged in a sequence from C4-dominant to C3-
dominant. It is not known if the snails in the region ingest
CAM plants. However, examined in the manner of Figure
8, it is evident that the y13C values of snail shells of the
southern Great Plains are generally indicative of the type of
vegetation in their immediate environment, although there
is still considerable scatter in the relationship. In the
regions where C4 plants were identified as the dominant
plant type, the transect-average y13C values of the snail
shell aragonite ranged from
4.3x to
1.9x with an
overall average of
2.8x. In localities where C3 plants
were documented to be the dominant type, these transect-
average values ranged from
10.1x to
8.8x with an
overall average of
9.0x (Fig. 8). In the areas with mixed
vegetation types, the shell y13C values are within the
extremes defined by the y13C values of the shells in areas
dominated by C3 or C4 plants. These observations are in
agreement with earlier observations (Francey, 1983; Good-
friend and Ellis, 2002; Goodfriend and Magaritz, 1987;
Metref et al., 2003; Stott, 2002).
22 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30

Table 1
Transect averages (all species) of measured shell y13C and y18O
Locality Transect Elevationa (m) Plant type Transect average Average y18O Model calculations
13 18
y C y O Calculated locality y18Ocalc D18O
Kubic 6 329 C3
8.5
2.1
2.0
2.2
0.2
Kubic 5 351 C4
3.5
1.9
Kubic 2 360 C4, C3
4.8
2.1
Kubic 4 360 C4
2.3
1.7
Kubic 3 366 C4
1.9
0.1
Bluff Creek 36 348 C3?
8.4
1.2
1.2
1.5
0.3
Salt Fork 9 320 C3
10.1
1.9
1.9
1.5 0.4
McDaniel 11 375 C4, C3
7.6
2.2
2.2
1.4 0.8
Burnham 17 519 C4, C3
8.3
0.8
1.0
0.9 0.1
Burnham 16 567 C4, CAM, C3
5.4
1.8
Burnham 12 607 C4, C3
5.8
2.3
Burnham 14 607 C4, CAM, C3
4.1
1.3
Burnham 13 610 C4
4.3
0.1
Big Salt Plain 15 482 C3
8.9
0.4
0.4
0.9
0.5
Skull Springs 18 671 C4, CAM, C3
6.0
1.6
1.6
0.3 1.3
Skull Springs 19 665 C4, C3
4.0
1.7
Hitch 22 915 C4, CAM
5.1
1.5
1.8
2.5
0.7
Hitch 21 933 C4, CAM, C3
5.0
0.6
Hitch 23 881 C3
8.5
2.3
Black Mesa 27 1312 C4, C3
6.1
2.7
2.2
2.1 0.1
Black Mesa 26 1324 C4, CAM, C3
6.6
1.5
Black Mesa 24 1488 C4, CAM
2.6
1.7
Black Mesa 25 1464 C4, CAM, C3
7.2
2.5
Black Mesa 35 1464 C4, CAM, C3
7.5
1.8
Owensby 61 2193 C3?
10.5
1.9
2.0
2.5
0.5
Owensby 60 2242 C3?
9.6
3.0
Owensby 59 2288 C3?
10.9
1.9
C.S. Ranch 30 1922 C4, CAM, C3
4.3
1.9
2.0
2.5
0.5
C.S. Ranch 29 1940 C4, CAM, C3
4.9
2.5
Chase 34 1952 C4, C3
9.2
3.8
2.5
2.5 0.0
Chase 33 2184 C4, C3
8.6
2.2
Chase 31 2220 C4, C3
9.0
2.5
Chase 32 2233 C4, C3
8.9
2.3
Locality averages of measured y18O. Also, y18O of shell at a locality as predicted from model calculations for diffusive evaporation (y18Ocalc). D18O =
y18Ocalc
y18Omeas.
a
Sea level datum.

For the transects representing the Owensby and Bluff
Creek localities, the plant species were not documented, but
the relatively negative land-snail shell y13C values suggest the
possible local dominance of C3 vegetation (hence the question
mark by the C3 symbol on the far right of the abscissa of Fig.
8). Some of the scatter in Figure 8 may be a result of the
complicating effects of incorporation of relatively 13C-rich
dietary carbonate from limestone (e.g., Goodfriend and Hood,
1983; Metref et al., 2003; Yates et al., 2002).
Oxygen isotopes
Average annual y18O of meteoric water is about
5.6x at
Norman, Oklahoma (USGS unpublished data, Martha
Scholl, personal communication), in the east and
9.8 x
at the higher elevations of Clovis, New Mexico (Nativ and
Riggio, 1990), in the west. As suggested by Figure 2, some of
this difference may be a consequence of differing proportions
of precipitation from different moisture sources and air
masses with different histories. Irrespective of the particular
mechanisms producing lower y18O values of average annual
precipitation at the higher western elevations, if the dominant
control on the y18O value of the snail shell aragonite was the
y18O value of annual precipitation, the shell y18O should be
lower at higher altitude. Figure 9a depicts snail shell y18O
values plotted against altitude for all of the analyzed
breplicationsQ (Appendix A). There is no correlation of shell
y18O with elevation evident in Figure. 9a.
Average y18O values of samples in each transect are listed
in Table 1 and plotted against elevation in Figure 9b. For the
transect-average values in Figure 9b, there may be a weak
relationship of shell y18O with altitude indicating some
tendency for a decrease of shell y18O with increasing
elevation. For an increase in elevation of ~2000 m, the slope
of the linear regression indicates a decrease in shell y18O of
only ~1x. However, even if this weak correlation in Figure
9b was significant, a decrease of ~1x is much less than the
decrease of ~4x expected if the y18O of annual precipitation
were the principal control on shell y18O values.
 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30 23

mentioned, these periods of activity are primarily evenings

and/or immediately after rainfall. From the archives of the
Climate Data Center of New Mexico State University
(www.weather.nmsu.edu), meteorological data for 1994
were available for one relevant station in New Mexico
(Clayton, see Appendix A). For the year 1994, hourly
meteorological parameters for seven stations in Oklahoma
(Appendix A) were available from Mesonet data (courtesy
of Oklahoma State University and University of Oklahoma).
Average temperatures characteristic of the aforementioned
conditions of land-snail activity appear to reasonably
represent the temperatures of the snail environment (Balak-
rishnan and Yapp, 2004), and these temperatures were
employed in our calculations (excluding days when temper-
atures were below 108C or above 278C). We also used
averages of nighttime RH for RH N 0.70 and 108C b T b
278C—i.e., the range conditions for snail activity. These
temperature and RH data are in Table 2. For the
calculations, it was assumed that essentially all of the water
imbibed by the snail was lost via evaporation (i.e., h = 0)
and that the ambient vapor was in isotopic equilibrium with
the input rain (for an explanation of model assumptions and
definition of terms, see Balakrishnan and Yapp, 2004).
Isotopic compositions of active season precipitation
were only available from three sites in the vicinity of the
study area: (1) Norman, Oklahoma (USGS unpublished
data, Martha Scholl, personal communication); (2) Ama-
rillo, Texas; and (3) Paducah, Texas (Nativ and Riggio,
1990). The average for Norman represents the years 1996–
Figure 7. y13C of southern Great Plains snail shell aragonite as a function of
elevation. (a) y13C values for all breplicationsQ; (b) average y13C values for
each transect (see text). Error bars in panel b represent one standard
deviation of the mean for the indicated transect.

The data of Figures 9a and 9b imply that factors other

than the y18O of annual precipitation are involved in
regulating the y18O values of the shells of land snails from
the southern Plains. In general, land snails are not active at
temperatures below 108C and above 278C (Cowie, 1984;
Thompson and Cheny, 1996) nor are they active at values of
relative humidity (RH) of less than about 0.70—expressing
RH as a decimal fraction (Van der Schalie and Getz, 1961,
1963). Thus, land snails are active only at night or following
rains (Cook, 1979; Edelstam and Palmer, 1950; Gelperin,
1974; Heatwole and Heatwole, 1978; Newell, 1966; Ward
and Slotow, 1992; Wells, 1944). Moreover, snail shells are
only precipitated when snails are active (Cowie, 1984).
Therefore, y18O values of snail shell aragonite should reflect
conditions within comparatively narrow ranges of high
relative humidities and moderate temperatures.
The steady-state flux balance model of Balakrishnan and
Yapp (2004) may provide some insight into the oxygen
isotope systematics of the snail shells of this study. The
Figure 8. Southern Great Plains land-snail shells. Open diamonds are
relevant model inputs for calculations of expected shell
transect-average y13C values of shells compared to the vegetation types at a
y18O values are temperatures, relative humidities (RH), and site (see text). Filled squares are average values of the respective transect
y18O values of precipitation estimated to be representative averages for each vegetation classification. Error bars are one standard
of the local environments at the times of snail activity. As deviation of the various means.
24 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30

Table 2
Active season temperature, relative humidity, and rainfall y18O
Weather station Summer T8C RHa y18O of summer rainb Rain isotope data station Proximal snail sample localities
Blackwell, OK 22.0 0.91
5.1 Norman, OK Kubic
Medford, OK 22.4 0.89
5.1 Norman Bluff Creek; Salt Fork
Cherokee, OK 22.0 0.89
5.1 Norman McDaniel
Alva, OK 21.2 0.88
5.1 Norman Burnham; Big Salt Plain
Beaver, OK 22.3 0.86
5.0 Paducah, TX Skull Springs
Goodwell, OK 22.8 0.87
6.7 Amarillo, TX Hitch
Boiser, OK 22.8 0.86
6.7 Amarillo Black Mesa
Clayton, NM 21.4 0.88
6.7 Amarillo Owensby; C.S. Ranch; Chase
a
RH as a decimal fraction.
b
See text for source of data.

2001, whereas the Amarillo data are for the years 1984/ and implies that the ambient relative humidity has an
1985. The average values are in Table 2. Model calcu- important influence on the y18O values observed in the shells
lations for each snail locality used the geographically (Balakrishnan and Yapp, 2004; Yapp, 1979). All other things
nearest active season meteorological data and isotopic being equal, the evaporation model predicts that a decimal
compositions of rain (Table 2). Because the measured fraction decrease in RH of only 0.01 produces a predicted
environmental parameters are not precisely coincident, increase in shell y18O of about 0.4x. This apparent
either spatially or temporally, with the respective snail sensitivity to RH and the westward decrease of 0.03–0.06
localities, some unknown error is introduced into the in the average active season nighttime RH (Table 2) may
comparisons of calculated and measured shell y18O values. partially compensate for the somewhat lower y18O values of
Nevertheless, the comparisons are instructive.
In the flux balance model of Balakrishnan and Yapp
(2004), it is assumed that the shell aragonite crystallized in
oxygen isotope equilibrium with snail body fluid that was
undergoing isotopic steady-state, diffusive evaporation. The
aragonite–water oxygen isotope fractionation equation of
Grossman and Ku (1986) is assumed to be applicable in
these model calculations. Let D18O = y18Ocalc
y18Omeas,
where y18Ocalc = the model-predicted y18O of the aragonite,
and y18Omeas = the measured y18O of the aragonite shell.
Note that D18O values of zero represent exact agreement
between predicted and measured y18O. For this compar-
ison, averages (Table 1) of measured y18O values of all
analyzed species at each locality were employed with the
idea that variations associated with differences among
individuals, species, times of shell formation, microenvir-
onments, etc., would be bsmoothed outQ and therefore
possibly better represent the average conditions reflected in
the meteorological data. These locality-average D18O
values, calculated with diffusive evaporation, scatter
around zero and with one exception differ from zero by
1x or less (solid diamonds in Fig. 10).
In contrast, for an assumption of oxygen isotopic
equilibrium between aragonite and land-snail body fluid
(local rain) that had experienced no evaporation prior to or
during snail activity, predicted shell y18O values are
significantly different from measured values. For this case,
the calculated D18O values differ from zero by more than
3.0x (solid triangles, Fig. 10), and all of these D18O values
for no evaporation are negative (
5.7 to
3.4x).
Figure 9. Southern Great Plains land-snail shells. Measured y18O values of
The fact that locality average D18O values for the diffusive
snail shell aragonite as a function of elevation. (a) y18O values of all
evaporation model scatter around and near zero suggests that breplicationsQ and (b) average y18O values for each transect (see text). The
this evaporation model may approximate the processes solid lines and associated equations in each figure represent the respective
operating in these land snails of the southern Great Plains linear regressions of the data.
 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30 25

Figure 10. D18O vs. average y18Omeas for each locality of southern Great Plains land snails. D18O = (y18Ocalc
y18Omeas). Diamonds represent the comparison
of calculated and measured shell y18O for shell y18O values calculated with diffusive evaporation of snail body fluid (solid diamonds: averages of all analyzed
specimens in a locality; shaded diamonds: averages of adult Vallonia only). Triangles represent D18O determined with the assumption that the snail shell was in
isotopic equilibrium with active season rain that experienced no evaporation either prior to or after being imbibed by the snails (solid triangles: averages of all
analyzed species in a locality; shaded triangles: averages of adult Vallonia only).

precipitation expected at the higher elevations and could Conclusion

explain why the shell y18O values are not well correlated with
elevation (Fig. 9).
Land snails of the genus Vallonia are commonly
associated with ancient sediments of the southern Great
Plains (Theler et al., 2004; Wyckoff et al., 1997), and the
diffusive evaporation model could provide a basis for
interpretation of many paleoenvironments in which this
genus was present. Therefore, y18O values of snail shell
aragonite predicted by the diffusive evaporation model
were compared only with the average measured y18O
values of adult Vallonia at each of the 12 modern
localities. With one exception, y18O values predicted by
the diffusive evaporation model (Balakrishnan and Yapp,
2004) differed from the measured y18O values of Vallonia
by no more than 0.8x (shaded diamonds, Fig. 10). At
present, we have no explanation for the single exception.
In contrast, for the case of no evaporation, all predicted
y18O values of Vallonia were significantly different from
measured y18O values (2.8–5.4x more negative, shaded
triangles, Fig. 10).
The approximate agreement between averages of meas-
ured shell y18O values of southern Great Plains land snails
and values predicted by the diffusive evaporation model is
in accord with the result obtained by Balakrishnan and Yapp
(2004), with reference to y18O data for western European
land snails measured by Lécolle (1985). Therefore, the
steady-state isotopic effects of evaporation (thus, relative
humidity) appear to be manifested in the y18O values of
land-snail shells of different species from two widely
separated regions with distinctly different climates.
At various southern Great Plains sample sites, transect-
average y13C values of land-snail shell aragonite are related
to the type of photosynthesis (i.e., C3, C4, or mixed) extant in
the local plant communities. There is considerable scatter in
the relationship, which suggests that caution should be
exercised in the interpretation of variations of shell y13C
values (e.g., Goodfriend and Hood, 1983; Metref et al., 2003;
Yates et al., 2002). However, measured y13C values of
coexisting Vallonia and Gastrocopta are well-correlated,
which appears to indicate similar feeding habits and suggests
that ancient samples of shells from these genera may be
useful sources of information on variations in southern Great
Plains plant ecology.
Measured y18O values of land-snail shells averaged over
these sample localities appear to be controlled primarily by
the local temperature, relative humidity, and y18O value of
rain at the times of snail activity (i.e., nighttime or after
rainfall in warmer months). This conclusion is supported by
the relatively good agreement between measured shell y18O
values and shell y18O values predicted with the evaporation
model of Balakrishnan and Yapp (2004).
Scatter of measured shell y18O values among and within
species at a site and among snails of different ages within a
genus (e.g., coexisting adults and juveniles of Vallonia)
indicates that the environmental information recorded by any
single small sample of land snails from the southern Great
Plains may depart significantly from the climatic bnormQ in a
locale. For paleoclimatic studies, such scatter emphasizes the
desirability of measuring, if possible, large numbers of
26 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30

individual snail shells from each defined sample to obtain Acknowledgments

average values that might better represent the larger
(bneighborhoodQ) scale temporal and/or spatial variations of
ancient climate that are of interest. The results presented here
for modern land-snail shells from the southern Great Plains of
North America suggest that meaningful climatic interpreta-
tions can be obtained from such averages, but it must be kept
in mind that the climatic information refers to the times of
snail activity and that more than one climatically relevant
parameter influences the shell y18O values.
We thank Martha Scholl of the United States Geological
Survey for data on the isotopic composition of precipitation
in Norman, Oklahoma, and Mesonet of the University of
Oklahoma and Oklahoma State University for meteorolog-
ical data. The paper benefited from the helpful comments of
D.D. Rousseau and an anonymous reviewer. This research
was supported, in part, by the Office of Graduate Studies at
SMU and NSF grant EAR-9614265 to C.J.Y.

Appendix A

Measured d13C and d18O values of Great Plains snails in this work

Locality Transect and replication Elevation (m) Plant type Land-snail species y13C y18O
Kubic 6B 329 C3 G. contracta
7.8
0.6
Kubic 6C 329 C3 G. contracta
9.1
2.2
Kubic 6C 329 C3 G. holzingeri
9.4
1.7
Kubic 6C 329 C3 G. pentodon
9.1
2.1
Kubic 6C 329 C3 V. parvula
8.9
3.1
Kubic 6C 329 C3 Vallonia sp. (juveniles)
8.8
3.1
Kubic 6C 329 C3 Glyphyalinia indentata
9.1
1.6
Kubic 6C 329 C3 Hawaiia minuscula
6.1
2.6
Kubic 5B 351 C4 G. procera
3.4
1.4
Kubic 5B 351 C4 V. parvula
2.9
2.3
Kubic 5B 351 C4 Vallonia sp. (juveniles)
3.7
2.7
Kubic 5C 351 C4 G. procera
3.4
1.2
Kubic 5C 351 C4 Vallonia sp. (juveniles)
4.3
1.9
Kubic 1B 351 C4 G. contracta
2.5
2.6
Kubic 1B 351 C4 G. procera
1.9
1.2
Kubic 1C 351 C4 G. procera
2.2
1.9
Kubic 2A 360 C4, C3 G. pellucida
4.3
2.5
Kubic 2A 360 C4, C3 G. procera
4.1
2.4
Kubic 2A 360 C4, C3 V. parvula
4.2
3.0
Kubic 2A 360 C4, C3 G. contracta
5.2
2.6
Kubic 2A 360 C4, C3 Vallonia sp. (juveniles)
5.4
0.9
Kubic 2A 360 C4, C3 Helicodiscus parallelus
5.7
3.4
Kubic 2B 360 C4, C3 G. armifera
6.1
2.3
Kubic 2B 360 C4, C3 G. holzingeri
7.1
2.1
Kubic 2B 360 C4, C3 G. pellucida
6.5
1.5
Kubic 2B 360 C4, C3 G. procera
6.4
1.8
Kubic 2B 360 C4, C3 Pupoides albilabris
2.1
1.7
Kubic 2B 360 C4, C3 V. parvula
4.9
2.5
Kubic 2B 360 C4, C3 Vallonia sp. (juveniles)
3.9
1.7
Kubic 2C 360 C4, C3 G. pellucida
3.9
1.7
Kubic 2C 360 C4, C3 G. procera
3.4
2.0
Kubic 2C 360 C4, C3 Vallonia sp. (juveniles)
2.8
2.9
Kubic 4A 360 C4 G. armifera
2.4
1.2
Kubic 4A 360 C4 G. contracta
0.7
2.8
Kubic 4A 360 C4 G. pellucida
2.4
2.2
Kubic 4A 360 C4 G. procera
3.1
2.5
Kubic 4B 360 C4 G. procera
2.7
2.1
Kubic 4C 360 C4 G. armifera
1.8
1.6
Kubic 4C 360 C4 G. contracta 0.0
1.6
Kubic 4C 360 C4 G. pellucida
1.6
1.7
Kubic 4C 360 C4 G. procera
3.1
0.4
Kubic 4C 360 C4 V. parvula
4.4
2.0
Kubic 4C 360 C4 Vallonia sp. (juveniles)
2.7
1.1
Kubic 3C 366 C4 G. procera
1.9
0.1
Bluff Creek 36A 348 C3? G. procera
6.8
1.1
 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30 27

Appendix A (continued )
Locality Transect and replication Elevation (m) Plant type Land-snail species y13C y18O
Bluff Creek 36B 348 C3? G. procera
10.0
1.3
Salt Fork 9A 320 C3 G. pellucida
10.3
1.2
Salt Fork 9A 320 C3 V. parvula
9.9
2.0
Salt Fork 9A 320 C3 Vallonia sp. (juveniles)
10.1
2.1
Salt Fork 9B 320 C3 G. pellucida
10.6
2.0
Salt Fork 9B 320 C3 V. parvula
10.0
2.1
Salt Fork 9B 320 C3 Vallonia sp. (juveniles)
10.0
1.8
Salt Fork 9C 320 C3 G. pellucida
10.0
1.7
Salt Fork 9C 320 C3 V. parvula
10.4
2.1
Salt Fork 9C 320 C3 Vallonia sp. (juveniles)
9.8
1.9
McDaniel 11C 375 C4, C3 G. procera
6.1
2.8
McDaniel 11A 375 C4, C3 G. procera
9.1
1.6
Burnham 17B 519 C4, C3 G. pellucida
9.4
0.8
Burnham 17B 519 C4, C3 G. procera
7.1
0.9
Burnham 16A 567 C4, CAM, C3 G. procera
4.8
1.9
Burnham 16B 567 C4, CAM, C3 G. procera
5.9
1.7
Burnham 12A 607 C4, C3 G. pellucida
5.4
2.3
Burnham 12A 607 C4, C3 G. procera
5.6
1.9
Burnham 12A 607 C4, C3 V. parvula
6.5
2.6
Burnham 14A 607 C4, CAM, C3 G. pellucida
2.8
2.0
Burnham 14A 607 C4, CAM, C3 G. procera
3.1
0.2
Burnham 14B 607 C4, CAM, C3 G. pellucida
5.9
0.7
Burnham 14B 607 C4, CAM, C3 G. procera
4.0
2.1
Burnham 14C 607 C4, CAM, C3 G. pellucida
4.9
0.9
Burnham 14C 607 C4, CAM, C3 G. procera
4.1
1.6
Burnham 13A 610 C4 G. pellucida
3.0 1.1
Burnham 13A 610 C4 G. procera
3.8 0.4
Burnham 13B 610 C4 G. pellucida
3.8 1.2
Burnham 13B 610 C4 G. procera
5.5
1.8
Burnham 13C 610 C4 G. procera
4.9
1.4
Big Salt Plain 15A 482 C3 G. pellucida
8.1
1.0
Big Salt Plain 15A 482 C3 G. procera
9.3
0.6
Big Salt Plain 15B 482 C3 G. procera
9.4 0.5
Skull Springs 18A 671 C4, CAM, C3 G. procera
5.1
2.2
Skull Springs 18A 671 C4, CAM, C3 P. albilabris
4.7
1.6
Skull Springs 18B 671 C4, CAM, C3 G. procera
6.8
1.0
Skull Springs 18C 671 C4, CAM, C3 G. procera
7.2
1.4
Skull Springs 19A 665 C4, C3 G. cristata
2.8
0.7
Skull Springs 19C 665 C4, C3 G. cristata
5.2
2.6
Hitch 22A 915 C4, CAM G. procera
4.2
1.5
Hitch 22A 915 C4, CAM P. albilabris
5.7
1.8
Hitch 22A 915 C4, CAM Succineidae
6.5
0.3
Hitch 22B 915 C4, CAM Succineidae
4.6
1.5
Hitch 22B 915 C4, CAM G. procera
4.4
2.2
Hitch 21A 933 C4, CAM, C3 G. procera
3.4
0.1
Hitch 21A 933 C4, CAM, C3 P. albilabris
5.6 0.0
Hitch 21C 933 C4, CAM, C3 Succineidae
5.9
1.6
Hitch 23A 881 C3 G. cristata
9.0
2.6
Hitch 23A 881 C3 G. procera
8.9
1.4
Hitch 23A 881 C3 P. albilabris
9.3
2.5
Hitch 23A 881 C3 V. parvula
9.9
2.7
Hitch 23A 881 C3 Vallonia sp. (juveniles)
9.4
3.2
Hitch 23B 881 C3 G. cristata
8.2
1.6
Hitch 23B 881 C3 V. parvula
8.8
2.2
Hitch 23B 881 C3 Vallonia sp. (juveniles)
8.9
2.8
Hitch 23C 881 C3 G. cristata
3.7
1.8
Black Mesa 27A 1312 C4, C3 V. gracilicosta
8.6
2.7
Black Mesa 27A 1312 C4, C3 Vallonia sp. (juveniles)
9.3
1.6
Black Mesa 27B 1312 C4, C3 G. procera
5.1
2.3
Black Mesa 27B 1312 C4, C3 V. gracilicosta
4.5
2.6
Black Mesa 27B 1312 C4, C3 Vallonia sp. (juveniles)
5.3
2.9
Black Mesa 27C 1312 C4, C3 Vallonia sp. (juveniles)
5.8
3.0
Black Mesa 27C 1312 C4, C3 G. pellucida
5.7
2.5
(continued on next page)
28 M. Balakrishnan et al. / Quaternary Research 63 (2005) 15–30

Appendix A (continued )
Locality Transect and replication Elevation (m) Plant type Land-snail species y13C y18O
Black Mesa 27C 1312 C4, C3 G. procera
6.2
2.5
Black Mesa 27C 1312 C4, C3 P. albilabris
3.6
2.4
Black Mesa 27C 1312 C4, C3 V. gracilicosta
6.3
3.1
Black Mesa 27C 1312 C4, C3 V. parvula
6.8
4.1
Black Mesa 26A 1324 C4, CAM, C3 V. gracilicosta
7.7
1.0
Black Mesa 26A 1324 C4, CAM, C3 Vallonia sp. (juveniles)
5.9
1.9
Black Mesa 26B 1324 C4, CAM, C3 G. pellucida
3.8
1.4
Black Mesa 26B 1324 C4, CAM, C3 G. procera
7.1
3.0
Black Mesa 26B 1324 C4, CAM, C3 V. gracilicosta
8.3
0.7
Black Mesa 26B 1324 C4, CAM, C3 Vallonia sp. (juveniles)
6.7
2.5
Black Mesa 24B 1488 C4, CAM P. albilabris
2.6
1.7
Black Mesa 25A 1464 C4, CAM, C3 G. pellucida
5.7
1.4
Black Mesa 25A 1464 C4, CAM, C3 G. procera
6.8
0.5
Black Mesa 25A 1464 C4, CAM, C3 V. gracilicosta
6.8
3.4
Black Mesa 25A 1464 C4, CAM, C3 Vallonia sp. (juveniles)
7.9
2.5
Black Mesa 25B 1464 C4, CAM, C3 G. pellucida
7.9
3.0
Black Mesa 25B 1464 C4, CAM, C3 V. gracilicosta
8.2
3.3
Black Mesa 25B 1464 C4, CAM, C3 Vallonia sp. (juveniles)
8.7
2.1
Black Mesa 25C 1464 C4, CAM, C3 G. pellucida
6.5
1.9
Black Mesa 25C 1464 C4, CAM, C3 V. gracilicosta
7.3
3.8
Black Mesa 25C 1464 C4, CAM, C3 Vallonia sp. (juveniles)
6.5
2.6
Black Mesa 35A 1464 C4, CAM, C3 G. pellucida
4.8
1.1
Black Mesa 35A 1464 C4, CAM, C3 V. gracilicosta
6.7
2.2
Black Mesa 35A 1464 C4, CAM, C3 Vallonia sp. (juveniles)
7.1
2.4
Black Mesa 35B 1464 C4, CAM, C3 G. pellucida
7.3
2.0
Black Mesa 35B 1464 C4, CAM, C3 V. gracilicosta
7.8
2.8
Black Mesa 35B 1464 C4, CAM, C3 Vallonia sp. (juveniles)
8.2
2.6
Black Mesa 35C 1464 C4, CAM, C3 G. pellucida
8.2
2.1
Black Mesa 35C 1464 C4, CAM, C3 Pupilla muscorum
8.7
1.7
Black Mesa 35C 1464 C4, CAM, C3 V. gracilicosta
8.4
0.8
Black Mesa 35C 1464 C4, CAM, C3 Vallonia sp. (juveniles)
8.1
0.5
Owensby 61A 2193 C3? P. muscorum
13.2
1.8
Owensby 61A 2193 C3? V. gracilicosta
11.1
1.7
Owensby 61A 2193 C3? Vallonia sp. (juveniles)
9.3
2.3
Owensby 61C 2193 C3? V. gracilicosta
8.4
1.6
Owensby 60B 2242 C3? Vallonia sp. (juveniles)
9.6
3.0
Owensby 59A 2288 C3? V. gracilicosta
10.4
3.1
Owensby 59A 2288 C3? Vallonia sp. (juveniles)
10.0
0.7
Owensby 59B 2288 C3? V. gracilicosta
10.9
1.3
Owensby 59B 2288 C3? Vallonia sp. (juveniles)
11.5
3.7
Owensby 59C 2288 C3? V. gracilicosta
11.7
1.0
C.S. Ranch 30A 1922 C4, CAM, C3 P. albilabris
4.3
1.9
C.S. Ranch 29B 1940 C4, CAM, C3 G. pellucida
4.7
2.4
C.S. Ranch 29B 1940 C4, CAM, C3 G. pellucida
5.0
2.6
C.S. Ranch 29C 1940 C4, CAM, C3 P. albilabris
5.0
1.0
Chase 34A 1952 C4, C3 V. gracilicosta
9.2
3.8
Chase 33A 2184 C4, C3 G. pilsbryana
6.9
3.3
Chase 33B 2184 C4, C3 Cionella lubrica
8.8
1.5
Chase 33B 2184 C4, C3 G. pilsbryana
9.3
2.5
Chase 33B 2184 C4, C3 G. pellucida
9.2
1.6
Chase 31C 2220 C4, C3 G. pilsbryana
8.5
1.3
Chase 31C 2220 C4, C3 Zonitoides arboreus
9.4
3.7
Chase 32C 2233 C4, C3 V. gracilicosta
9.3
1.8
Chase 32C 2233 C4, C3 Vallonia sp. (juveniles)
8.6
2.7

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