
(browse) and cool growing season or aquatic grasses, whereas
C
4
plants include most tropical and temperate grasses”
(MacFadden et al., 1999). The difference in the ratios of
13
C
to
12
C in these two categories of plants is related to the fact that
they have different photosynthetic pathways. C
3
plants, which
use the Calvin Cycle as their photosynthetic pathway, concen-
trate less
13
C relative to
12
C than is the case with C
4
plants,
which use the Hatch-Slack photosynthetic pathway.
In mammalian herbivores, there is a consistent difference in
the
13
Cto
12
C ratio of the tooth enamel between those that pri-
marily feed on C
3
plants (browsers) and those that primarily
feed on C
4
plants (grazers). Those that are mixed feeders have
intermediate ratios. However, the values measured in the tooth
enamel are not exactly the same as in the plants themselves.
Rather, it has been established empirically that in all cases the
tooth enamel is enriched in
13
C relative to the plants on which
the animal feeds. The degree of this enrichment appears to be
size dependent, with mouse-sized animals having one frac-
tionation value and larger animals having a different, larger
fractionation value. However, despite this variation, modern
herbivores that feed exclusively on C
3
plants can be distin-
guished from those animals that feed solely on C
4
plants by ana-
lyzing their
13
C/
12
C ratio in tooth enamel. Mixed feeders range
in between the C
3
and C
4
endpoints. Making the assumption
that this difference was also the case with fossil mammalian her-
bivores is the basis for interpreting diets of the extinct forms.
Using carbon isotope data together with wear facet analysis
from the teeth of six different 5 million year old horses from
Bone Valley Florida, MacFadden et al. (1999) were able to
reconstruct their dietary preferences; ranging from browsers to
mixed feeders to grazers. Wear facet analysis corroborated the
isotopic results. This was surprising because on the basis of
the high crowned nature of the horse teeth alone, they all would
have been considered grazers. This result is concordant with
paleoenvironmental, “...reconstructions of central Florida at
5 Ma [which] indicate low-elevation floodplain and estuarine
environments with a mosaic of local close-canopy forests, wood-
lands, and open-country grasslands” (MacFadden et al., 1999).
In this instance, isotopic ratios of vertebrate fossils were utilized
to corroborate a paleoenvironmental reconstruction made from
other lines of evidence and to caution against the use of a single
indicator of dietary preference, such as crown height.
The other most common isotope system to be investigated
in fossil vertebrates is that of oxygen, despite the complicat-
ing factors mentioned above. The ratio of
18
Oto
16
O in verte-
brates is sensitive to temperature, humidity and diet (Kohn and
Cerling, 2002). In one recent study using fossil taxa, measure-
ments of the oxygen isotope ratio in both terrestrial mammals
and gar in the Big Horn Basin of Wyoming were interpreted as
indicating that a rapid increase in mean annual temperature
occurred at the end of the Paleocene and beginning of the Eocene
(Fricke et al., 1998). This study circumvented the problem of var-
iation in diet affecting the oxygen isotope ratio by sampling the
same genera through the period of time under investigation.
Because mammalian molars generally grow from the tip of
the cusps to the base of the roots, by measuring the ratio of
18
Oto
16
O at different heights of the enamel of a horse molar
it was possible for Bryant et al. (1996) to observe the
18
O/
16
O ratio through the time that the tooth formed instead
of taking an average for the entire tooth. The changes they
observed were regarded as due to seasonal temperature changes
through that period. Because molars develop consecutively,
with the first molar forming, then the second, and finally
the third molar, it was possible for Bryant et al. (1996)to
correlate between molars and estimate the seasonal fluctua-
tions in temperature over the entire period that formation of
the molars took place. An excellent, more technical summary
of this topic is summarized in Parrish (1998, pp. 84–88
and 154–161).
Thomas H. Rich and Patricia Vickers-Rich
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Cross-references
Carbon Isotopes, Stable
Cretaceous Warm Climates
Isotope Fractionation
Nearest-Living-Relative Method
Oxygen Isotopes
Paleotemperatures and Proxy Reconstructions
Pleistocene Climates
ANIMAL PROXIES, VERTEBRATES 15