Earth and Planetary Science Letters, 24 (1974) 91-98 [~]

©North-Holland Publishing Company, Amsterdam - Printed in The Netherlands

180 A N D 13C IN T H E S H E L L S O F F R E S H W A T E R M O L L U S C S A N D
THEIR ENVIRONMENTS

P. FRITZ and S. POPLAWSKI

Department o f Earth Sciences, University of Waterloo, Waterloo, Ont. (Canada)

Received May 21, 1974

Revised version received July 20, 1974

Carbon and oxygen isotope analyses on shells of freshwater molluscs and their habitat are presented. The data ob-
tained reconfirm the usefulness of such lao analyses for paleoenvironmental and paleohydrologicalstudies.
The laC analyses on freshwater molluscs from lakes in southwestern Ontario, specimensgrown under laboratory
conditions and a comparison with the tSC contents of the dissolved inorganic carbon in their habitat show that the
l~C contents in mollusc shells are primarily co~rolled by the aqueous carbonate species. Vital effects and food con-
trol appear to have only minor importance. The significance of this observation on the usufulness of mollusc shells for
14C dating is discussed.

1. Introductory remarks Much less significance for this type of research

Following the development of double collecting
mass-spectrometers for precise stable isotope analyses
a number of researchers focused their attention on
the 180 fractionations in the carbonate-water system
[ 1 - 3 ] . These investigations revealed that many
marine molluscs deposit their shells in isotopic equili-
brium with the water of their habitat and the deter-
mination of 180/260 ratios in fossil shells therefore
provided a tool for paleotemperature studies. In order
to calculate the actual temperature of ancient environ-
ments it is necessary, however, to know the oxygen
isotopic composition of the water in which these
molluscs lived. This prerequisite can often be evaluated
for marine environments but it is difficult to predict
the 2aO contents of ancient freshwater bodies. Never-
theless, it has been recognized that the 2aO contents
of freshwater molluscs are useful paleoenvironmental
indicators since also their shells are formed in isotopic
equilibrium with the water [4]. This is especially true
if approximate growth temperatures are known since
in this case the isotopic composition of the water body
can be determined. Such information is useful not
only in paleoenvironmental investigations but is ex-
tremely valuable for paleohydrological studies which
attempt to elucidate the history of freshwater systems.
has been attached to the 13C contents of mollusc shells
although there is good evidence that some species
deposit their shells in isotopic equilibrium with the
aqueous carbonate species of their habitat [5]. In
general the impression persists that the carbon uptake
by molluscs is rather complicated especially since it
was found that modern shells can have 14C concentra-
tions which give 14C ages in excess of 2000 years [6,7].
At present most lSC analyses on mollusc shells are
done in conjunction with 14Cdeterminations. The fact
that mollusc shells could give false age data is extremely
important and several studies were under taken to ex-
plain the reasons for this observation and to find pos-
sible means for correcting these data. The general con-
clusion from these studies was, that the uptake of tac-
free organic food or limestone particles was primarily
responsible for the 14C deficiencies in these shells. This
interpretation was supported by growth experiments
during which carbonate regenerated under controlled
conditions on damaged shells was analyzed [9]. Ad-
ditional suggestions emphasized possible biological
fractionations and the importance of aqueous carbon-
ate species [4-6,8] which would be more significant
for the abundance of 13C in the mollusc shells than for
14C concentrations [7].
Because of this rather confusing picture it was felt
92
that additional growth experiments might provide data
which could explain some of these abnormal variations.
In this paper the 13Cresults obtained on molluscs
grown under controlled conditions are presented and
compared with data obtained from modern specimens
in a number of small lakes in southwestern Ontario
(Canada), as well as some observations made on ma-
terial separated from cores obtained in one of these
lakes.
The experimental work differs from previous studies
inasmuch as care was taken not to damage the shells
and hereby upset the physiology of the animals, and
to utilize, where ever possible, only molluscs which
were bred in the laboratory. It was speculated that
biological fractionations may well be associated with
physiological changes although at present we do not
have any proof for this assumption [10].
No ~4C determinations have been made since not
yet enough carbonate for 14C analyses has been gen-
erated in our laboratory. It is anticipated, however,
that this will be achieved in the near future and these
results will be published at a later date.
2. Experimental and analytical techniques
The molluscs analyzed for this study were collected
in several small lakes in southwestern Ontario, others
were separated from cores of lake sediments or were
grown in the laboratory under controlled conditions.
The lake samples include both live and dead speci-
mens. Water samples were collected for ~So and ~3C
analyses in the habitat of these molluscs. Since this
sampling was mostly done in the shallow, near-shore
environments the isotope data may not be representa-
tive for the lakes as a whole.
The dissolved inorganic carbon in the tank experi-
ments was artificially controlled by bubbling CO2-free
air and 13C-labelled CO2 through the tank water. The
tank bottom was covered with carbonate-free sand
but some finely powdered calcite had to be added
periodically in order to maintain a near-neutral pH in
the water. Care was taken to prevent the snails from
coming in contact with this carbonate and ingesting
it with their food. The carbonate and the macroscopic
food (lettuce) were identical in all experiments, but
no attempt was made to control algae gi'owth which
provided the foor for some of the species. The evapo-
P. FRITZ AND S. POPLAWSKI
rated water in these tanks was replenished continuous-
ly via a simple overflow mechanism and only deion-
ized water was added.
For the isotope analyses of the mollusc shells or-
ganic matter was removed from the crushed shells
by treatment with 5% sodium hypochlorite solution
for 6 - 1 2 hours. The samples were then thoroughly
washed with distilled water and treated under vacuum
with 100% H3PO4 [3]. The evolved CO: was separated,
purified and analyzed for 180 and 13Cin a double
collecting mass-spectrometer.
The 180 analyses of the water were done on CO2
equilibrated at 25°C with an aliquot of the water
sample [11 ]. The X3C contents of the total dissolved
inorganic carbon in lake and tank water were measured
as CO2 removed from the water sample by acid treat-
ment under vacuum.
All isotope date are expressed in the conventional
6%onotation. They refer to the standard PDB for all
~3C analyses and the lSo contents of mollusc shells.
The IaO concentrations of the water samples are re-
ported with respect to the standard SMOW. The ana-
lytical precision of the 180 and 13C data for both
water and carbonate samples was better than -+0.2%o.
3. Results and discussion
3.1. 180 in lakes and freshwater molluscs
The 180 contents of smaller lakes is dependent on
the 180 contents of the inflowing water and the local
precipitations but is equally strongly influenced by
evaporation and exchange reactions with the atmos-
pheric vapor reservoir. The relationship between
surface area and depth of the lake is thereby of great
importance. Most shallow lakes show pronounced
seasonal variations in their 180 contens with the
highest values observed during the summer months.
Little or no seasonal variations are observed in large,
deep lakes but climatic changes over long periods
will cause a response in the stable isotope contents.
In northern climates most lakes are ice covered
during the winter months and inflowing surface water
and groundwater during this period cause a decrease
in the 180 concentrations. The lowest 8180 values are
normally observed immediately following the spring
melt and the heavy inflow of isotopically light waters.
With the onset of the ice-free season strong 180 en-
180 AND ~3C IN THE SHELLS OF FRESHWATER MOLLUSCS 93

richments will occur in small lakes due to the prefer-
ential loss of H2 ~60 by evaporation. The highest ~So
concentrations throughout the year will be observed
during the months of August to October.
The maximum level of ~aO enrichment is not only
controlled by the climatic parameters which control
the rate of evaporation and thus the isotope fractiona-
tions which occur, but also by the degree of flushing
which is related to the amounts of surface and sub-
surface inflow and outflow from these lakes. Each
lake develops a very distinct pattern of ~So variations
throughout the year which is only indirectly related
to the average annual precipitation. However, preci-
pitation has a significant indirect influence since it
controls the 180 contents of the inflowing waters.
Major climatic changes can thus easily introduce
changes in the average ~So contents of lake waters and
the associated seasonal variations. The significance of
climatic changes has been documented not only for
small lakes [4] but is also very pronounced in larger
systems such as the Great Lakes [ 12].
The importance of the hydrodynamic regime of
small lakes is emphasized in the data shown in Fig. 1.
Each of the four lakes considered shows a different
pattern of seasonal variations in its 180 contents and
the different levels of ~80 enrichment during the
summer months are clearly visible. The most stagnant
lakes show the highest ~80 contents whereas lower
8180 values were found in lakes with significant sur-
face and subsurface inflow and outflow.
Fig. 1 also demonstrates that most mollusc species
closely reflect the 180 contents of these lakes during
the growth season. Differences in ~80 contents of
various species from the same lake are almost cer-
tainly related to different growth periods and the
temperature of their habitat (shallow vs. deep water).
However, the overall variations between different
species are small, an observation confirmed by data
obtained from a core drilled into the sediments of a
small lake in this area (Fig. 2) [13], and samples ob-
tained from a core of Lake Erie sediments [12]. These
small differences which rarely exceed 2%0 are not sur-

8'%*/0. 2A)"
-4-

-6"
SPRY LAKE tl° 2A2-

2A3-
o+emo

~,-©Z~o

-8- 2BI'

2B2" o qlt~ z~
- I0 - - FRANCIS LAKE
~1./11 i f /
2B$-
/
-12- 3AI'
1975
A M J J A S 0 3A2" +e,~ •

\/ /
8'~0%. 3A5'
-4-
5A4" © IH- eL5
LITTLE LAKE

3A5- • ,~em o
\ \ \ \
5BI' •,5,o

5B2"
-I0-
~///~AT LAK~E -~o -fo -~o -~o -4o 4o 3 0 -~o -io 6 .io .~o
~'80 PDB%° ~3C PDB%o
• P,s,#,#m mlJd~m contortu~
-12-
1975 ~OP , s , ~ m v~lnnco=dm
4, Pl~d~m feftugtn~m ¢osfotum
A M d d A' S ' 0 ~ V o / ~ t a m c o r , ~ #~confuso
• Hel/so~ co~totum mpch/g~$e
• Amntcola / ~ t o n t
Fig. 1. ~So contents in waters from small lakes in southwestern
Ontario, Canada, and the average values and range of ~80 con- Fig. 2. 180 and 13Cconcentrations in mollusc shells separated
tents in modern mollusc shells collected in these rakes. Note from a core drilled in the sediments of Townline Lake (south-
that the water data are expressed as 8 ~80%o SMOW and the western Ontario, Canada). The total length of the core is
shell data as 8 lSO%o PDB. about 2 m and possibly covers several thousand years [ 13].
94
prising because the increasing water temperatures du-
ring the growth season which would cause lower180
contents in the carbonate deposited are at least in
part compensated or even overcompensated by the in-
crease in 180 in the water due to evaporation. These
data thus demonstrate again that it is virtually im-
possible to deduce temperature information from 180
analyses on freshwater molluscs but that such analyses
can be useful in paleoenvironmental or paleohydrolog-
ical studies especially if a number of different species
can be compared.
3. 2 Carbon-13
3. 2.1. General remarks
Whereas there is general agreement that the 180
contents in shells of freshwater molluscs are good en-
vironmental indicators there is considerable doubt
as to the usefulness of carbon isotope investigations.
This is due to several facts: The laC variations which
may occur in freshwater systems were only studied
in a very limited number of investigations and our
knowledge at present is very incomplete. Furthermore
14C data demonstrate that molluscs can derive some
of the carbon used in the shell carbonate from 14C de-
ficient sources. These may be either the organic food
[6,8], inorganic carbonate utilized in the gizzard to
mechanically disintegrate plant tissues [ 14] or inor-
ganic dissolved carbon [4,5,14]. This dissolved carbon
may be of biogenic origin, be derived from the disso-
lution of carbonates or be isotopically controlled by
exchange with the atmospheric carbon dioxide reser-
voir.
In a attempt to solve the question on the origin of
the carbon in mollusc shells we grew molluscs in 13C-
controlled environments. As will be shown, the data
obtained from this study indicate that the inorganic
dissolved carbon closely controls the 13C contents
of the carbonate shells deposited. A brief discussion
of the 13C in inorganic dissolved carbon in lake waters
is therefore warranted.
3.22. Carbon-13 o f lake waters
The most comprehensive investigation of 13C in
dissolved inorganic carbon in lake waters was presented
by Oana and Deevey [15]. They discussed in detail
the various sources and their influence on the 13C con-
tents of the different dissolved carbon species and
P. FRITZ AND S. POPLAWSKI
showed that most of the observed variations in the
lakes they investigated can be explained by exchange
with the atmospheric carbon dioxide reservoir (513C =
- 7 to - 9 [16,17]), the 13C content of the inflowing
groundwater (dissolved carbonates) and CO2 produced
during the decomposition of organic matter. The con-
tributions from these different sources results in these
lakes in a 13C stratification. Surface waters usually
have the highest 813C values which typically fall be-
tween - 5 and -9%owhereas deeper water is strongly
influenced by biological activities. The aerobic decom-
position of organic matter and plant material normal-
ly results in the production of CO2 with 813C values
below -20%0. If anaerobic conditions prevail, reduced
carbon species, such as methane, are generated simul-
taneously to the production of CO2. Laboratory ex-
periments [18] and field data [15] show that under
these conditions CO2 may become considerably en-
riched in 13C with respect to the original plant ma-
terial. Mixing of such CO2 with other dissolved carbon
present typically produces 6 laC values in deeper waters
between - 1 0 and -20%0.
Another process which may significantly influence
microenvironments within a lake is the photosynthetic
activity of aquatic plants. Similar to terrestrial plants,
aquatic plants utilize preferentially isotopically light
carbon which results in a 13C enrichment in the im-
mediate vicinity of these plants. For example, calcium
carbonate precipitated on pond weeds is enriched in
13C with respect to the aqueous carbonate as a direct
consequence of the preferential 12C extraction by the
pond weeds [14,19].
The effects make it impossible to predict the 13C
contents of the dissolved inorganic carbon in a lake.
However, similar to the 1So contents of lake water,
one might expect that each lake is characterized by
specific 13C concentrations in its dissolved carbon
species. Furthermore, these concentrations are to a
large degree influenced by biological activities, which
in these latitudes depend on the climatic conditions
prevailing and therefore 13C may still prove to be a
useful paleoenvironmental indicator especially if con-
sidered together with 180 data. One could thus spec-
ulate that a combined consideration of 180, laC and
lac in present day lakes and their geologic environ-
ment could help in the identification of lakes in which
the contribution of 14C-free carbon is minimal and
which also in the past produced organic matter or
180 AND 13C IN THE SHELLS OF FRESHWATER MOLLUSCS 95

O" TABLE 1
)leo%* SMOW-H20 I
13C contents of mollusc shells and dissolved inorganic carbon
-2-
from tank experiments
-4.
Species Sam- Average Range
p j o ples 613C
-6
ana-
-8'
lyzed

-I0 Tank 1 aqueous CO 2 5 --13.1 --11.3 to --14.4

-12
%/o Ly mnaea
stagnalis 8 -13.8 -13.1 to-14.3
o
Helisoma
o,o, I trivolve 3 -14.3 -13.7 to -15.2
-iz -b -~ -~ -a -~ o z 4 6 Viviparus 6 -14.2 -11.1 t o - 1 4 . 4
Fig. 3. The 180 contents of lake waters is compared with the Sphaerium
t3C contents of their total dissolved inorganic carbon. The ar- lacustre 16 -14.2 mixed samples
rows indicate the isotopic evolution during spring and summer. Tank 4 aqueousCO2 4 -35.5 -35.7 to -35.3
Lymnaea
stagnalis 2 -31.7 -30.5 to -33.0
shells which can be used for 14C dating purposes. The Helisoma
stable isotopes would be used as a m o n i t o r for the past trivolve 1 -33.4
history of the lake. Viviparus 1 -32.9
It is noteworthy that a comparison o f 180 and ~3C Tank 5 aqueousCO2 3 + 5.4 + 6.0 to + 4.9
data obtained from samples collected at different Lymnaea
times o f the year in a number o f lakes indicate a posi- stagnalis 7 + 0.8 - 0.3 to+ 1.3
tive correlation between the abundances o f the two Helisoma
trivolve 1 + 5.3
isotopes in water and dissolved carbonate (Fig. 3). A
possible explanation for this 180-13c relationship
could be that shallow and more stagnant lakes (high
180 contents) have higher rates o f biological activities tance [4,8]. In order to investigate in more detail the
and both the anaerobic decomposition o f their b o t t o m importance o f the dissolved inorganic carbon which,
mud as well as the photosynthetis activities o f aquatic as shown above, is directly related to the environment
plants contribute isotopically heavy CO2 to the lake in which the molluscs live we grew several species
water. Furthermore exchange reactions with atmos- o f molluscs in tanks with artificially controlled 13C
pheric CO2 might be more significant in shallow lakes contents in the dissolved carbon. The results of
than in deeper ones, which also would contribute these studies are given in Table 1 and, in part, are
laC-rich dissolved inorganic carbon. As the latter pro- shown in Fig. 4. Fig. 4 also compares the 13C con-
cess would add modern carbon to the water it is of tents of the total dissolved inorganic carbon of several
special importance for 14C studies. Note, however, lakes with the 13C content o f their mollusc fauna.
that our samples were always collected close to shore, The seasonal variations in 13C concentration in these
within the habitat o f the molluscs collected, and these lakes are shown in Fig. 5 and compared to the average
results are therefore not necessarily representative for and range o f 613C values of shells from a number of
these lakes as a whole. mollusc species.
An inspection o f these data indicates a surprisingly
3.2.3. Carbon-13 in freshwater molluscs good correlation between the ~3C contents of mollusc
All previous investigators emphasized the importance shells and total dissolved inorganic carbon. Since in
o f environmental parameters to explain the 13C con- the tank experiments the food source was kept as con-
tents in freshwater mollusc shells. Vital effect such as stant as possible and the tank b o t t o m s consisted of
biological fractionations which could occur during the carbonate-free sand, these results strongly suggest
shell deposition were thought to be o f minor impor- that primarily the dissolved inorganic carbon and not
96 P. FRITZ AND S. POPLAWSKI

HO
~mC%* PDB- MOLLUSCA
SPRYTANKLA'K~E5 ~ /
*5
• Lym~ooo Sh~F~h:S
• ................ LIT;::L:A::KE ]~ / , / ~ ,~
0
:J;"2~21,-,%°''°° FRANCISLAKE
®_~ !
~'~C % . PDB
-5'
o& ÷4-

-I0.
÷2- SPRY
~BOATLAKE
-15"
0
-20"
-2
-25- LITTLE LAKE
.....

-50-

-~5- ~'~C %0 PDB - CO~q

-35 -30 -z5 -20 -~5 -,0 -5 0 .5 .Jo

Fig. 4. A comparison of 13C contents of mollusc shells and

the total dissolved inorganic carbon of their habitat. The in-
-I0- ~-
organic carbon data are average values of the aqueous carbon-
ate species during the growth season.
-12- ~

the food source control the '3C contents of mollusc

shells. Note that the species analyzed include gastro-
pods as well as pelecypods. These results do not agree
with one of the assumptions made by Keith et al. [8]
which attempted to explain abnormal ~4C values in
modern molluscs as being due to the uptake of ~4C-free
organic food. However, these authors also suggest that
carbon dioxide from humus decay might be involved
in the shell formation. This agrees with our observa-
tions and supports the findings of Digby [20] who
indicated that the carbon in the shell carbonate of
Mytilus edulis is derived by direct precipitation of
aqueous carbonate. Furthermore, since the isotope
fractionation between bicarbonate, aqueous carbon-
ate and solid carbonate is small, these data indicate
that the shell carbonate is deposited at least in near
equilibrium with the dissolved bicarbonate or carbon-
ate. This interpretation agrees with the results of Mook
and Vogels [5].
Minor carbon contributions from the food source,
however, could account for the observation that in
high 13C environments (tank 5 and lakes) the shell
carbonates have somewhat less 13C than the dissolved
carbon, and in the low ~3C environment (tank 4) the
shells are somewhat richer in ~3C than the dissolved
carbon. We have not determined the ~3C concentration
1975
A M d d A S 0
Fig. 5. Seasonal variations of the 13C contents of lakes in
southwestern Ontario and the average values and range
of 13C contents in modern mollusc shells from these lakes.
in the food material, but it can be assumed that it con-
tains more t3C than the dissolved carbon in tank 4 and
less than the one in tank 5 [ 16,21 ].
As mentioned above, not all species grown in our
tanks fed on the lettuce provided. For example the
sphaeriids (pelecypods) are filter feeders and live in
the sediment, Viviparus fed almost exclusively on algae
and only Lymnaea and Helisoma ingested both lettuce
and algae. Fortunately, the 13C contents in algae are not
a simple function of 13C contents of the dissolved
carbon dioxide but depend also in a non-linear fashion
on the partial pressure of the dissolved carbon dioxide
[21 ]. In our experiments the ~3C contents of the feed
gas was kept constant but we did not attempt to control
the partial pressure of the CO2 in the tanks and would
therefore expect very significant 13C differences be-
tween different algae generations and between algae
grown in the different tanks. These differences are
not apparent in the shell material analyzed which
1So AND taC IN THE SHELLS OF FRESHWATER MOLLUSCS 97

• Lymnaeo stagnalis These differences are not only a feature of these

A Galba elodes
• He//soma trivolve
(D Helisoma campanula/urn
0 Helisoma anceps
• Sphaerium simile
• • • ~
• • ~
A • CI 0 CHESLY LAKE
• •
modern lakes but were found equally consistent
throughout the length of the core from Townline
Lake (Fig. 2) and in samples from a Lake Erie core
which covers a time span exceeding 10,000 years.
Such consistency is difficult to explain, and possibly
reflects the minor micro-environmental, biological or
food influences. Perhaps the artificial environment
created in our tanks and the uniform food available
SPRY LAKE
prevented us from detecting these effects. If it is at
present impossible to document the relative impor-
GOULD LAKE tance of vital affects and food control it should be
possible to distinguish between the two if more and
better controlled tank experiments are carried out
and if stable isotope analyses are compared with 14C
data.
~ O LITTLE LAKE

A • ~ BOAT LAKE 4. Conclusions

A • t[D FRANCIS LAKE
A • EUGENIA LAKE
A' C o
Fig. 6. Differences in 13C contents in the shells of different
mollusc species from various lakes in southwestern Ontario,
Canada.
again supports the argument that only a minor food
control exists.
Each mollusc species lives in a rather well-defined
micro-environment. We did not investigate in detail
these micro-environments since often only bulk sam-
piing of shells washed to the shorelines provided enough
well-preserved material for our study. If, however, the
613C values of several species found in different lakes
are compared the importance of these micro-environ-
ments can be documented. Fig. 6 is such a comparison
which shows the surprisingly constant pattern of
613C values: Lymnaea stagnalis and Galba sp. are in-
variably the species with the lowest 13C concentrations
whereas Helisoma anceps shows the highest 813C values.
The overall spread in 613C values for samples from
the same lake is almost certainly related to differences
in these micro-environments
With this study an attempt is made to explain the
carbon isotopic composition of freshwater mollusc
shells and to reopen the discussion on the usefulness
of 14C dates obtained on such material. Experimental
data and data obtained on shells collected in small
lakes in southwestern Ontario indicate that the 13C
contents in mollusc shells are primarily controlled by
the dissolved inorganic carbon and its XSc contents.
Vital effects and food controls appear to be minimal
though it cannot be excluded that they have some
influence. We hope to substantiate these findings with
14C analyses.
The 'aC contents of the aqueous carbonate species
in a lake depend on a variety of factors which cannot
be evaluated a priori. It is possible, however, to use
'4C analyses on modern mollusc shells and the aqueous
carbonate of their habitat to identify lakes in which
at present the contributions of 14C-free carbon is min-
imal. A comparison of 180 and 13C data from mod-
ern samples and shells separated from the sediments
of such lakes could then possibly provide a tool to es-
timate or evaluate such contributions in the past and
to select shell material on which reliable 14C dates
may be obtained.
A comparison of 180 contents in lake waters with
those of the carbonate shells show that the numerical
values of the 8 lSO-SMOW data of the water samples
collected during the summer months are close to those
98
o f the 5180-PDB o f the mollusc shells. This agreement
demonstrates again the usefulness o f 180 analyses on
mollusc shells in paleoenvironmental or paleohydrolog-
ical investigations.
Acknowledgements
The authors wish to acknowledge the technical
assistance given by Mr. R. Drimmie and Mr. D. Killey.
The manuscript greatly benefited from l o n g discussions
with J.C. Fontes, University o f Paris. The Townline
Lake core was collected by Dr. T.W. A n d e r s o n , CCIW,
Burlington, w h o described in detail the p a l y n o l o g y
o f this core [13]. The drafting has been done by
Mr. P. Russell. Financial support was given through
N.R.C. Grant A. 7954.
References
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a paleotemperature scale, J. Chem. Phys. 18 (1950) 849.
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Carbonate-water isotopic temperature scale, Geol. Soc.
Am. Bull. 62 (1951) 417.
3 S.E. Epstein, R. Buchsbaum, H.A, Lowenstam and
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4 M. Stuiver, Oxygen and carbon isotope ratios of fresh-
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5 M.G. Mook and J.C. Vogel, Isotopic equilibrium between
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P. FRITZ AND S. POPLAWSKI
oxygen isotopic composition of mollusk shells from ma-
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