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Isotopic discrimination of resource partitioning among ungulates in C3-

dominated communities from the Miocene of Florida and California

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DOI: 10.1666/05006.1

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Paleobiology, 32(2), 2006, pp. 191–205

Isotopic discrimination of resource partitioning among

ungulates in C3-dominated communities from the Miocene of
Florida and California

Robert S. Feranec and Bruce J. MacFadden

Abstract.—Stable isotope analysis of mammalian tooth enamel is a valuable method for examining
resource partitioning in modern and ancient environments where there is a mixture of C3 and C4
plants. However, before 7 Ma North American ecosystems were composed predominantly of C3
plants, complicating isotopic assessment of resource partitioning. Study of modern African and
North American ecosystems has shown that niche partitioning among mammals may be discerned
in communities dominated by C3 plants, suggesting that a similar approach may work for ancient
C3 ecosystems. Here, such analyses are applied to explore resource use and niche partitioning in
two ancient C3-dominated communities, one from California and one from Florida. Each locality,
Black Hawk Ranch (California) and the Love Bone Bed (Florida), occurs in Miocene deposits that
accumulated prior to the rapid increase in C4 ecosystems 7 Myr ago. d13C and d18O values were
obtained from the tooth enamel of eight species from Black Hawk Ranch, and 15 species from the
Love Bone Bed. Results from the 197 bulk isotope samples showed significant differences in d13C
among taxa at the Love Bone Bed, but no significant differences were observed among taxa at Black
Hawk Ranch. At both localities, equids generally have more positive d13C values than co-occurring
taxa, suggesting that equids occupied more open habitats, whereas antilocaprids, camelids, and
proboscideans have more negative values, implying utilization of more closed communities. One
result of note is the positive d13C values of Pediomeryx (Yumaceras) hamiltoni from the Love Bone
Bed, which suggests that P. (Y.) hamiltoni incorporated abundant fiber, possibly grass, in the diet
similar to the horses from this locality. The lack of significant differences among taxa at Black Hawk
Ranch may indicate a relatively homogeneous flora, or presence of abundant resources permitting
niche overlap, whereas the opposite is implied by the presence of significantly different isotope
values among taxa at the Love Bone Bed. The results from this study highlight the utility of isotopic
techniques allowing discernment of resource partitioning in C3-dominated landscapes such as
those that persisted for the millions of years before the rapid increase in C4 ecosystems that oc-
curred during the late Miocene.

Robert S. Feranec.* 3060 Valley Life Sciences Building, Department of Integrative Biology, University of
California, Berkeley, California 94720
Bruce J. MacFadden. Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611.
E-mail: bmacfadd@flmnh.ufl.edu
* Present address: Department of Biological Sciences, Herrin Hall, Stanford University, Stanford, Califor-
nia 94305. E-mail: feranec@stanford.edu

Accepted: 4 November 2005

Introduction enamel is a valuable method for examining re-

Assessment of resource use and niche par-
titioning among taxa through space and time
has important implications for interpreting
community structure and dynamics. For ex-
ample, if these attributes can be followed
through time, it becomes possible to test clas-
sic assumptions about communities, such as
Hutchinson’s (1958) proposition that where re-
sources are limiting, ecologically similar taxa
must partition resources to ensure persis-
tence, or that resource overlap is generally
avoided by partitioning diet, habitat, or time
of use (Schoener 1974).
Stable isotope analysis of mammalian tooth
q 2006 The Paleontological Society. All rights reserved.
source partitioning in modern and ancient en-
vironments where there is a mixture of C3 and
C4 plants. (DeNiro and Epstein 1978; Vogel
1978; Quade et al. 1992; Wang et al. 1994;
Bocherens et al. 1996; MacFadden and Cerling
1996; Cerling et al. 1997a; Koch et al. 1998).
Aspects of resource partitioning revealed by
stable isotope analysis include diet and habi-
tat use, as well as temporal partitioning of re-
sources (Koch et al. 1989; Cerling and Sharp
1996; Fricke and O’Neil 1996; Jahren et al.
1998; Balasse et al. 2003). However, isotopic
discrimination of resource partitioning has
rarely been shown in C3-dominated commu-
0094-8373/06/3202-0003/$1.00
192 ROBERT S. FERANEC AND BRUCE J. MAC FADDEN
FIGURE 1. Location of Miocene fossil sites used in this study. Fauna at both locations were deposited about 9.5 Ma
during the Clarendonian land mammal age.
nities even though prior to 7 Ma North Amer-
ican communities were composed predomi-
nantly of C3 plant taxa (Cerling et al. 1993,
1997b; Ehleringer and Monson 1993). Meth-
odologies to analyze resource use in mam-
malian taxa before the rapid expansion of C4
plants ca. 7 Ma include comparison to modern
analogs, correlation to various morphological
characteristics, and community body size
structure (cenograms), which are able to sug-
gest diet and vegetation structure (Legendre
1986; Janis 1988, 1990; Janis and Ehrhardt
1988; Solounias et al. 1995; Fortelius and So-
lounias 2000; Fortelius et al. 2002). Isotopic
analysis of modern ecosystems from Africa,
Europe, and North America has shown that
discerning niche partitioning among mam-
mals is possible in communities dominated by
C3 plants (Bocherens 2003; Feranec 2003, 2004;
Cerling et al. 2004).
This study examines resource use among
ungulates from two North American fossil lo-
calities that sample communities prior to the
rapid increase in C4 ecosystems (Cerling et al.
1993, 1997b; Ehleringer and Monson 1993). We
address the following questions: (1) Can sta-
ble isotope values from large samples of an-
cient mammals reveal resource partitioning in
C3-dominated paleoenvironments in North
America? (2) If resource partitioning is dis-
cernable, do the results corroborate what is
currently inferred about ancient taxa through
prior analysis such as examination of mor-
phology? (3) Are there general patterns in the
isotopic values observed among taxa that per-
sist across the continent? This technique may
provide another method to independently as-
sess ecology in ancient taxa and determine if
these results concur with the results of other
methodologies during the extensive amount
of time prior to 7 Ma.
Background
Fossil Localities. The Love Bone Bed (LBB)
locality of Alachua Co., Florida (Fig. 1), has
been assigned biochronologically to the late
Clarendonian, confined to the latest Claren-
donian (Cl3) land mammal age (Webb et al.
1981; Tedford et al. 2004). At least 84 species
of marine and terrestrial vertebrates have
been identified at this locality, including over
40 mammal species. Webb et al. (1981) suggest
that the terrestrial vertebrates of LBB come
from three different habitats—a riparian hab-
itat, a forest, and more open habitat.
Contemporaneous with LBB is the Black
Hawk Ranch (UCMP V3310) locality of north-
ern California, which is located (Fig. 1) on the
southern side of Mount Diablo in Contra Cos-
ta Co., California (Prothero and Tedford 2000).
This locality has been placed biochronologi-
cally into Cl3 (Tedford et al. 2004) and has
 RESOURCE PARTITIONING IN MIOCENE COMMUNITIES
been assigned to Chron C4Ar, dated to be-
tween 9.7 and 9.0 Ma (Prothero and Tedford
2000). At least 20 mammalian species are
known from Black Hawk Ranch (BHR), and
plants identified from quarries near the mam-
mal locality have been suggested to be indic-
ative of floodplain and woodland-chaparral-
type habitats (Condit 1938; Axelrod 1944).
Isotopes in Paleoecology. This study exam-
ines the d13C and d18O values in the tooth
enamel of ancient mammals. Isotopic results
are expressed in standard d-notation: X 5
[(Rsample/Rstandard) 2 1] 3 1000, where X is the
d13C or d18O value, and R 5 13C/12C and 18O/
16
O, respectively. In this study, the d13C and
d O values are reported relative to the V-PDB
18
standard.
Carbon Isotopes in Mammalian Diet. Tissues
of animals that feed on different kinds of
plants will reflect the carbon isotope value of
their food (DeNiro and Epstein 1978; Tieszen
1978; Vogel 1978; Quade et al. 1992). Carbon
isotope values are useful in diet studies be-
cause the three different photosynthetic path-
ways used by plants, C3, C4, and crassulacean
acid metabolism (CAM), impart different 13C/
12
C ratios to plant tissues. Tropical, warm-
growing-season grasses and sedges using the
C4, or Hatch-Slack, photosynthetic pathway
are relatively enriched in the heavy carbon
isotope (13C). These plants have a mean d13C
value of 213.0‰ and generally range from
29‰ to 219‰ (O’Leary 1988; Farquhar et al.
1989; Ehleringer et al. 1991; Ehleringer and
Monson 1993). On the other hand, C3, or Cal-
vin cycle, plants including most trees and
shrubs, as well as grasses in regions with cool
growing seasons, are relatively enriched in the
light carbon isotope (12C). C3 plants have a
mean d13C value of 227.0‰ and typically
range from 222‰ to 235‰ (O’Leary 1988;
Farquhar et al. 1989; Ehleringer et al. 1991; Eh-
leringer and Monson 1993). The third path-
way, the CAM pathway, is characteristic of
succulents (e.g., cacti) and may yield values
that range between the end members for C3
and C4 plants (O’Leary 1988; Ehleringer et al.
1991; Ehleringer and Monson 1993). CAM
photosynthesis is typically found in more re-
stricted xeric habitats, CAM plants are gener-
ally not widespread on the landscape (Ehler-
193
inger et al. 1991), and very few extant large
ungulates use succulents as a significant part
of their diet (Janis and Ehrhardt 1988). There-
fore, it is assumed that herbivores generally
did not make succulents a significant part of
their diet, and the isotopic effects of CAM
photosynthesis are only considered briefly in
this study.
Studies utilizing differences in the carbon
isotope values from mammals generally focus
on communities containing a mixture of C3
and C4 plants, thus enabling taxa to be distin-
guished on the basis of the predominant for-
age included in the diet (Vogel 1978; Tieszen
1994; MacFadden and Cerling 1996; Cerling et
al. 1997a, Koch et al. 1998; MacFadden et al.
1999). Taxa that feed on tropical grasses and
sedges are considered to be grazers, those
taxa that feed on trees and shrubs are brows-
ers, and taxa that feed on a mixture of tropical
grasses and sedges and trees and shrubs are
considered intermediate feeders (sensu Hof-
mann and Stewart 1973). These studies are re-
stricted in their scope because C4 plants are to-
day confined to lower latitudes; at higher lat-
itudes (above 438N in North America) plants
almost entirely utilize the C3 photosynthetic
pathway (Teeri and Stowe 1976; Stowe and
Teeri 1978; Tieszen et al. 1979; Tieszen et al.
1997; Sage et al. 1999). It also appears that al-
though C4 plants may have been present in
ecosystems throughout the Miocene, especial-
ly in the Great Plains (Fox and Koch 2003),
most herbivorous taxa, particularly herbivo-
rous taxa in Florida, restricted their diets to C3
plants until about 7 Ma (MacFadden and Cer-
ling 1996; Cerling et al. 1997b). Different pro-
cesses, however, such as variation in light in-
tensity, temperature, nutrient availability, and
water stress, can produce variation in the d13C
value in C3 plants (Farquhar et al. 1989;
O’Leary et al. 1992; Ehleringer and Monson
1993; Koch 1998). C3 plants typically have
more negative values in closed, forested hab-
itats, whereas in more open, drier habitats
more positive isotope values are characteristic
(Farquhar et al. 1989; van der Merwe and Me-
dina 1991; O’Leary et al. 1992; Ehleringer and
Monson 1993; Koch 1998; Cerling and Harris
1999; Cerling et al. 2004). Recent studies have
used these habitat differences to highlight the
194 ROBERT S. FERANEC AND BRUCE J. MAC FADDEN
use of carbon isotopes in discriminating re-
source partitioning of herbivorous taxa in
pure C3-dominated environments (Katzen-
berg 1989; Quade et al. 1995; Bocherens et al.
1997; Cerling et al. 1997a; Cerling and Harris
1999; Katzenberg and Weber 1999; Iacumin et
al. 2000; Bocherens 2003; Drucker et al. 2003;
Richards and Hedges 2003; Cerling et al. 2004;
Drucker and Bocherens 2004; MacFadden and
Higgins 2004).
Stable Carbon Isotope Values in Paleoecology.
Herbivores reflect the carbon isotope values of
plants ingested, but the d13C value of the tooth
enamel is further enriched by a consistent
amount, 114.1 6 0.5‰ for medium- to large-
bodied mammals (Cerling and Harris 1999).
Setting the fractionation from plant material
to tooth enamel at 114.1‰ in large mamma-
lian herbivores, extant taxa that feed solely on
C3 plants will display enamel carbon isotope
values between 220.9‰ and 27.9‰. Modern
taxa that feed solely on C4 plants exhibit iso-
topic values around 0.0‰ (Quade et al. 1992;
Cerling et al. 1993; Koch 1998). The carbon in
plants and ultimately enamel is derived from
the atmosphere. The d13C value of atmospheric
CO2 has decreased 21.5‰, from 26.5‰ to
about 28.0‰, because of fossil fuel burning
over the last 200 years (Friedli et al. 1986; Ma-
rino and McElroy 1991; Marino et al. 1992).
Data further suggest that Miocene atmospher-
ic CO2 d13C values were more positive than
pre-industrial levels (Pagani 1999; Passey et al.
2002). For this reason, enamel isotope values
from ancient taxa are expected to be about
1.5‰ 6 1‰ more positive than the modern
taxa, owing to d13C differences in the atmo-
sphere during the past as compared to the
modern (Koch et al. 1998; Passey et al. 2002).
For fossil taxa then, a diet of pure C3 plants
would range from 220.4‰ to 26.4‰. Isotope
values more positive than 26.4‰ would im-
ply incorporation of either C4 or CAM plants
into the diet. The 26.4‰ d13C value as indic-
ative of C4 or CAM plant incorporation into an
herbivore diet appears conservative, as previ-
ous studies on modern and fossil ungulates
show that pure C3 feeders rarely have values
more positive than 28.0‰ (MacFadden and
Cerling 1996; Cerling and Harris 1999; Cer-
ling et al. 2004).
Oxygen Isotopes in Tooth Enamel. The oxy-
gen isotope composition in mammalian tooth
enamel is dependent on the isotopic compo-
sition of ingested water, the consistent frac-
tionation of oxygen isotopes between body
water and the tooth enamel, and the metabo-
lism of the particular animal (Land et al. 1980;
Longinelli 1984; Luz et al. 1984; Luz and Ko-
lodny 1985; Koch et al. 1989; Kohn 1996; Kohn
et al. 1996). Mammalian herbivores ingest wa-
ter from two sources, either through drinking
meteoric water or from consumed plants. Me-
teoric water is affected by climatic influences
such as temperature and humidity, such that
d18O values are more positive where and when
it is warmer and more negative where and
when it is colder (Dansgaard 1964; Rozanski
et al. 1992; Fricke and O’Neil 1999; Kendall
and Coplen 2001; Kohn and Welker 2005).
However, being located within the same gen-
eral area at the same time, the isotopic values
of ingested meteoric waters are assumed to be
similar at a particular fossil locality. Water in
plant leaves also varies and is typically more
positive than the local meteoric waters as a re-
sult of evapotranspiration. Enrichment in d18O
values in the leaves is more pronounced in
warmer and more arid conditions (Yakir et al.
1990; Yakir 1992; Ometto et al. 2005). Thus, an-
imals that occupy open habitats would be ex-
pected to ingest more positive d18O as com-
pared to taxa foraging in a cooler, moister for-
ested habitat.
Body size and individual metabolism have
a large impact on the oxygen isotope compo-
sition of tooth enamel. In mammals, large, ob-
ligate-drinking taxa with a low metabolism
are suggested as the most likely to accurately
reflect the d18O ingested (Longinelli 1984; Luz
et al. 1984; Bryant and Froelich 1995). All taxa
analyzed at the two fossil localities are large
and considered to have a lower metabolism.
Because it is not possible to know whether the
studied taxa were obligate drinkers or to
know the metabolism of each individual, the
d18O values are scrutinized with caution.
Terminology for Habitats. In this study, we
refer to habitats as being ‘‘open’’ versus
‘‘closed’’ rather than providing specific habi-
tat types such as woodland or grassland.
Open habitats generally refer to grasslands
 RESOURCE PARTITIONING IN MIOCENE COMMUNITIES
FIGURE 2. Predicted representation of isotopic data in
C3-dominated environments. We expect taxa inhabiting
closed environments to display relatively negative car-
bon and oxygen isotope values, whereas we expect open
environment inhabitants to display more positive val-
ues.
and savannas, whereas closed habitats refer to
woodlands and forests. This is because al-
though more positive d13C values are observed
in open habitats and more negative d13C val-
ues in closed habitats (Fig. 2), there is not a
strict relationship between the percentage of
tree cover and stable carbon isotope value in
any habitat (van der Merwe and Medina 1991;
Magnusson et al. 2002; Rossetti et al. 2004).
Materials and Methods
Preparation of Tooth Enamel Samples. Tooth
enamel of a total of 138 specimens from 15
species were analyzed from the Love Bone
Bed of central Florida. This includes data from
ten specimens from MacFadden and Cerling
(1996). Taxa include two camelids, ‘‘Hemiauch-
enia’’ minima and Procamelus grandis, one dro-
momerycid, Pediomeryx (Yumaceras) hamiltoni,
three equinine horses, Calippus cerasinus, Cal-
ippus elachistus, and Protohippus gidleyi, five
hipparionine horses, Cormohipparion ingen-
uum, Cormohipparion plicatile, Nannippus wes-
toni, Neohipparion trampasense, and Pseudhip-
parion skinneri, two rhinocerotids, Aphelops ma-
lacorhinus and Teleoceras proterum, one gom-
photheriid, Gomphotherium sp., and one
tapirid, Tapirus simpsoni.
Fifty-nine specimens of eight species were
sampled from the Black Hawk Ranch (BHR)
locality of the Clarendonian land mammal
195
age of northern California. The taxa sampled
from BHR include one antilocaprid, Caprome-
ryx sp., two camelids, Megatylopus sp. and Pro-
camelus sp., one oreodontid, Merychyus major,
one tayassuid, Prosthenops sp., two equids,
Hipparion forcei and ‘‘Pliohippus’’ leardi, and
one gomphotheriid, Gomphotherium productum.
The BHR fauna is contemporaneous in age
with the LBB fauna, ranging from 8 to 10 Ma
(Tedford et al. 2004).
Preference was given to sampling the pre-
molars and the third molar because these
teeth are among the last ones to develop, min-
eralize, and erupt in ungulates (Severinghaus
1949; Fuller 1959; Rees et al. 1966; Wegrzyn
and Serwatka 1984; Hillson 1986), and our
goal was to recognize the resource partition-
ing in adult animals. Where the premolars
and third molars were few, these were sup-
plemented by sampling first or second molars
or incisors.
The general method for stable isotope sam-
pling of tooth enamel followed MacFadden
and Cerling (1996) and Koch et al. (1997). For
the tooth enamel, sampling involved drilling
20–30 mg of pristine enamel powder off the
tooth along a non-occlusal surface parallel to
the growth axis and across its entire length us-
ing a 0.5 mm inverted cone carbide drill bit
and a variable-speed Dremely rotary tool.
The powder was first collected and treated
with 30% hydrogen peroxide for 24 hours to
remove organics. The hydrogen peroxide was
then decanted and the powder was then
washed with distilled water, and soaked in 0.1
N acetic acid for another 24 hours to remove
any diagenetic carbonate. The following day it
was again decanted and washed with distilled
water, rinsed with 100% ethyl alcohol, and
dried overnight. Although ontogenetic sam-
pling was not completed on any of the teeth
in this study, the height of the teeth and the
width of enamel in the analyzed taxa indicate
that the sampled segment of tooth represents
a significant portion of at least one year (Kohn
2004).
After treatment, the samples were analyzed
with an ISOCARB automated carbonate prep-
aration system attached to a Micromass Op-
tima gas source mass spectrometer within the
Geology Department at the University of Cal-
196 ROBERT S. FERANEC AND BRUCE J. MAC FADDEN

TABLE 1. Stable carbon and oxygen isotope values from taxa of the Love Bone Bed, Florida.

Mean d13C d13C SD Mean d18O d18O SD

Taxon n (‰) (‰) (‰) (‰)
Love Bone Bed total 138 211.6 1.3 21.3 2.7
Camelidae
(A) ‘‘Hemiauchenia’’ minima 10 213.3 0.7 22.4 0.9
(B) Procamelus grandis 11 212.2 0.6 22.5 1.6
Dromomerycidae
(C) Pediomeryx (Yumaceras) hamiltoni 11 210.5 0.8 22.7 1.0
Equidae
(D) Calippus cerasinus 2 210.9 0.2 2.9 0.3
(E) Calippus elachistus 2 210.2 0.7 2.9 2.8
(F) Cormohipparion ingenuum 13 210.8 0.8 21.7 2.2
(G) Cormohipparion plicatile 12 211.4 0.5 21.5 2.2
(H) Nannippus westoni 11 211.2 0.8 21.1 3.0
(I) Neohipparion trampasense 11 210.8 0.8 20.5 3.6
(J) Protohippus gidleyi 10 211.2 0.7 22.1 2.3
(K) Pseudhipparion skinneri 13 29.9 1.3 0.0 4.5
Gomphotheriidae
(L) Gomphotherium sp. 9 212.6 0.6 21.2 2.9
Rhinocerotidae
(M) Aphelops malacorhinus 5 212.8 0.3 0.5 0.8
(N) Teleoceras proterum 5 213.3 0.4 1.1 0.4
Tapiridae
(O) Tapirus simpsoni 13 213.2 0.7 21.8 2.1

ifornia, Davis. The ;1-mg samples were dis-

solved in 100% phosphoric acid at 908C to cre-
ate CO2.
Mean differences among species within lo-
calities were compared by ANOVA and post
hoc LSD tests, which are similar to t-tests but
take into account multiple comparisons. Sta-
tistical analyses were run on JMP IN 5 for stu-
dents, with significance set at p , 0.05. Pre-
cision was 0.1‰ for d13C and 0.1‰ for d18O.

FIGURE 3. Stable carbon and oxygen isotope values
from taxa at the Love Bone Bed, Florida. In comparison
to Black Hawk Ranch (Fig. 4), the LBB species show sta-
tistically significant differences and a wider range in
mean d13C values. Symbols represent the mean, dotted
lines represent one standard deviation around the mean.
Symbols: square, Camelidae; triangle, Dromomerycidae;
circle, Equidae; plus sign, Gomphotheriidae; hexagon,
Rhinocerotidae; diamond, Tapiridae. Abbreviations: A,
‘‘Hemiauchenia’’ minima; B, Procamelus grandis; C, Pediom-
eryx (Yumaceras) hamiltoni; D, Calippus cerasinus; E, Cal-
ippus elachistus; F, Cormohipparion ingenuum; G, Cormo-
hipparion plicatile; H, Nannippus westoni; I, Neohipparion
trampasense; J, Protohippus gidleyi; K, Pseudhipparion skin-
neri; L, Gomphotherium sp.; M, Aphelops malacorhinus; N,
Teleoceras proterum; O, Tapirus simpsoni.
Results
Carbon and Oxygen Isotopic Values from Love
Bone Bed, Florida. The 138 samples from the
Love Bone Bed display a mean d13C value at
211.6‰ 6 1.3‰ (6 SD) (Table 1, Fig. 3; see Ap-
pendix online at http://dx.doi.org/10.1666/
05006.s1), and show statistically significant
differences in their d13C values (Table 2). It ap-
pears from Figure 3 that the taxa can be split
into two distinct groups based on their carbon
isotope values, those taxa that are more posi-
tive than the locality mean and those taxa
with more negative mean values. The group of
taxa more positive than the locality mean in-
clude all eight horse species as well as the dro-
 RESOURCE PARTITIONING IN MIOCENE COMMUNITIES 197

TABLE 2. Significant differences in d13C value among taxa from the Love Bone Bed, Florida. Abbreviations: y, sig-
nificant difference observed at p , 0.05; Y, significant difference observed at p , 0.01; Y, significant difference
observed at p , 0.001; -, no significant difference observed.

momerycid Pediomeryx (Y.) hamiltonii. Within
this group the equid Pseudhipparion skinneri
has the most positive mean d13C value at
29.9‰ 6 1.3‰ as well as the widest range in
values, from 211.3‰ to 27.8‰ for 13 sam-
ples. The taxon displaying the most negative
mean d13C value is Cormohipparion plicatile,
which has a mean carbon isotope value of
211.4‰ 6 0.5‰ with a range from 212.5‰
to 210.8‰ for 12 samples. The group of taxa
that have d13C values more negative than the
locality mean include two camelid species,
two rhinocerotids, a tapirid, and a probosci-
dean. Of this group, the taxon displaying the
most positive mean d13C value is Procamelus
grandis, which has a mean value of 212.2‰ 6
0.6‰ with a range from 213.2‰ to 210.8‰
for 11 specimens. Teleoceras proterum, with a
mean d13C value of 213.3‰ 6 0.4‰ and a
range from 213.7‰ to 212.8‰ for five spec-
imens, has the most negative mean value
within this grouping. There is a statistically
significant correlation (p , 0.001) between the
number of samples and the range of carbon
isotope values observed within a taxon. Sim-
ilar to the results from BHR (see below), there
is no trend in the isotopic data when compar-
ing hypsodont taxa with brachydont taxa.
The mean d18O value for the 138 samples at
LBB is 21.3‰ 6 2.7‰, and there are no sta-
198 ROBERT S. FERANEC AND BRUCE J. MAC FADDEN

TABLE 3. Stable carbon and oxygen isotope values from taxa of Black Hawk Ranch, California. The sampled gom-
phothere specimens were fragmentary and assigned only to the Gomphotheriidae; but as only Gomphotherium prod-
uctum is known from this locality, these specimens are here referred to G. productum? in the table.

Mean d13C d13C SD Mean d18O d18O SD

Taxon n (‰) (‰) (‰) (‰)
Black Hawk Ranch total 59 211.8 0.7 25.0 1.3
Antilocapridae
(A) Capromeryx sp. 10 211.8 0.9 24.1 1.3
Camelidae
(B) Megatylopus sp. 8 211.4 0.6 25.1 1.0
(C) Procamelus sp. 11 211.8 0.7 25.1 0.7
Equidae
(D) Hipparion forcei 9 212.1 0.4 26.1 0.7
(E) ‘‘Pliohippus’’ leardi 5 211.6 0.4 25.1 1.4
Gomphotheriidae
(F) Gomphotherium productum? 7 212.2 0.5 25.8 1.4
Oreodontidae
(G) Merychyus major 4 211.7 1.1 22.9 1.5
Tayassuidae
(H) Prosthenops sp. 5 211.6 0.4 25.3 0.2

tistically significant differences among taxa
(Table 1, Fig. 3). The two equinine species of
Calippus have the most positive d18O values. C.
cerasinus has a mean d18O value of 2.9‰ 6
0.3‰ with a range from 2.7‰ to 3.1‰ for two
FIGURE 4. Stable carbon and oxygen isotope values
from taxa at the Black Hawk Ranch, California. Symbols
represent the mean; dotted lines represent one standard
deviation around the mean. The mean d13C values of
BHR species group together, compared with the species
from the Love Bone Bed (Fig. 3), which are more spread
out. Symbols: open circle, Antilocapridae; closed square,
Camelidae; closed circle, Equidae; plus sign, Gomphoth-
eriidae; times sign, Oreodontidae; open square, Tayas-
suidae. Abbreviations: A, Capromeryx sp.; B, Megatylopus
sp.; C, Procamelus grandis; D, Hipparion forcei; E, ‘‘Plio-
hippus’’ leardi; F, Gomphotherium productum?; G, Mery-
chyus major; H, Prosthenops sp.
specimens, whereas C. elachistus has a mean at
2.9‰ 6 2.8‰ with a range from 0.9‰ to
4.8‰ for two specimens. The most negative
mean d18O value is displayed by Pediomeryx
(Y.) hamiltonii at 22.7‰ 6 1.0‰ with a range
from 24.7‰ to 21.3‰ for 11 specimens.
There is no statistically significant correlation
between the number of samples and the range
in oxygen isotope values.
Carbon and Oxygen Isotopic Values from Black
Hawk Ranch, California. The d13C values for
the 59 samples from Black Hawk Ranch are
relatively constrained around a mean of
21.8‰ 6 0.7‰ (Table 3, Fig. 4, Appendix).
This mean is similar to that of LBB. However,
in contrast to LBB, there are no statistically
significant differences in d13C values among
any of the eight taxa examined. The camelid
Megatylopus sp. has the most positive mean
d13C value at 211.4‰ 6 0.6‰ SD with a range
in values from 212.2‰ to 210.6‰ for eight
specimens. The proboscidean taxon sampled,
Gomphotherium productum, has the most nega-
tive mean d13C value at 212.2 6 0.5‰ with a
range in values from 213.0‰ to 211.7‰ for
seven specimens. There is no significant cor-
relation in the range of carbon isotope values
and the number of samples taken per taxon (p
. 0.24). Further there does not appear to be
 RESOURCE PARTITIONING IN MIOCENE COMMUNITIES
any trend in the isotopic data comparing hyp-
sodont taxa such as the horses with brachy-
dont taxa such as the peccary and oreodont.
The range of d18O values among taxa is
much wider than the range seen in the d13C re-
sults for the BHR specimens (Table 3, Fig. 4).
The mean d18O for the BHR samples is 25.0‰
6 1.3‰. Also in contrast to the d13C values,
there are significant differences among taxa.
The horse Hipparion forcei has the most nega-
tive mean d18O value at 26.1‰ 6 0.7‰ and a
range from 27.2‰ to 24.8‰ for nine speci-
mens. The values for H. forcei are similar to
those for all taxa except Capromeryx sp. and
Meychyus major. The d18O values for ‘‘P.’’ leardi
are not different from those of any of the other
sampled taxa at BHR. The proboscidean, Gom-
photherium productum, has the widest range in
d18O values, with a mean value of 25.8‰ 6
1.4‰ and a range from 27.8‰ to 23.5‰. G.
productum is similar to all taxa except M. major
and Capromeryx sp. Capromeryx sp. is statisti-
cally different from all taxa except G. produc-
tum and H. forcei. The two camelids, Megaty-
lopus sp. and Procamelus grandis, have similar
d18O values. Both camel taxa are similar to
each of the other sampled taxa except M. ma-
jor. The peccary, Prosthenops sp., has the small-
est range in isotope values (25.7‰ to 25.1‰)
and is significantly different from Capromeryx
sp., ‘‘Pliohippus’’ leardi, and M. major. M. major
has the most positive d18O value with a mean
at 22.9‰ 6 1.5‰ and a range from 24.9‰
to 21.6‰ for four specimens. The oreodonts
were significantly different from all taxa ex-
cept Capromeryx sp. and ‘‘P.’’ leardi.
Discussion
Love Bone Bed. The statistically significant
differences observed among taxa suggest dif-
ferent diets, use of different habitats, or both.
Webb et al. (1981) noted that the LBB taxa are
derived from at least three terrestrial habitat
types: riparian, forested, and open-habitat.
Although we expect that it will be difficult to
distinguish riparian from forested habitats
isotopically, comparing the isotopic values ob-
tained with the patterns observed among taxa
in both modern and ancient environments
(Bocherens 2003; Feranec 2003, 2004; Cerling
et al. 2004) suggests that more positive d13C
199
and d18O values likely indicate residence in
more open habitats, whereas more negative
isotope values indicate habitation of more
closed environments.
The high diversity of horses (presence of at
least nine species) represents a unique oppor-
tunity to examine isotopically resource use
prior to the rapid increase in C4 ecosystems
that occurred ca. 7 Ma. Coming from the same
clade and having relatively similar morphol-
ogies, isotopic differences should represent
true resource use differences rather than the
possibility of clade-level variation in isotope
incorporation. Because significant differences
in d13C values are observed among the equids,
taxa do appear to be partitioning resources
(Table 2, Fig. 3). The more positive d13C values
in C. cerasinus, C. elachistus, and C. plicatile like-
ly indicate use of open habitat, a conclusion
further supported by elevated d18O values.
The more negative oxygen isotope values ob-
served in the other horses, such as C. ingenuum
and P. gidleyi, may indicate foraging in a more
closed environment compared to the other
equids, such as using woodland or the ecotone
between open and forested habitat. Because of
issues related to the d18O source, metabolic in-
fluences, and the fact that there are no signif-
icant differences in d18O among taxa at LBB, it
is not possible to ascribe particular habitat use
based on the oxygen isotope values, even
though we might presume that the positive
18
O values may indicate less dependence on lo-
cal meteoric waters and a higher dependence
on leaf water (leaf water generally being more
enriched relative to meteoric waters) (Helliker
and Ehleringer 2000; Yakir and Sternberg
2000; Ometto et al. 2005).
The isotopic values for the remaining taxa,
except Pediomeryx (Y.) hamiltoni and ‘‘Hem-
iauchenia’’ minima, corroborate previously sug-
gested niche use (Table 4) (Webb et al. 1981;
MacFadden 1998), such that the negative car-
bon and oxygen isotope values observed in
Gomphotherium sp. and Teleoceras simpsoni in-
dicate residence in a closed environment,
whereas the intermediate d13C values of Pro-
camelus grandis probably represent use of both
open and forested habitats. Further discrimi-
nation as to whether the closed environment
was riparian is not currently possible using
200 ROBERT S. FERANEC AND BRUCE J. MAC FADDEN

TABLE 4. Range in d13C values and environmental interpretation of isotopic values for the Love Bone Bed. Abbre-
viations: Y, supports previously suggested habitat for taxon; N, does not support previously suggested habitat for
taxon; ?, not enough data to distinguish between habitat types.

Supported by
Taxon d13C range (‰) Suggested habitat d13C values
Camelidae
‘‘Hemiauchenia’’ minima 214.6‰ to 212.3‰ open habitat N
Procamelus grandis 213.2‰ to 210.8‰ open habitat ?
Dromomerycidae
Pediomeryx (Yumaceras) hamiltoni 211.9‰ to 29.0‰ forest N
Equidae
Calippus cerasinus 211.0‰ to 210.7‰ open habitat Y
Calippus elachistus 210.7‰ to 29.7‰ open habitat Y
Cormohipparion ingenuum 212.3‰ to 29.0‰ open habitat Y
Cormohipparion plicatile 212.5‰ to 210.8‰ open habitat Y
Nannippus westoni 212.0‰ to 29.2‰ open habitat Y
Neohipparion trampasense 212.0‰ to 29.5‰ open habitat Y
Protohippus gidleyi 212.3‰ to 210.4‰ open habitat Y
Pseudhipparion skinneri 211.3‰ to 27.8‰ open habitat Y
Gomphotheriidae
Gomphotherium sp. 213.7‰ to 211.7‰ riparian Y
Rhinocerotidae
Aphelops malacorhinus 213.2‰ to 212.4‰ open habitat ?
Teleoceras proterum 213.8‰ to 212.8‰ open habitat/riparian* ?
Tapiridae
Tapirus simpsoni 214.3‰ to 211.7‰ forest Y
* T. proterum suggested as a riparian inhabitant on basis of morphology in Webb et al. (1981), whereas MacFadden (1998) suggested that T. proterum
is an open-country inhabitant based on basis of stable isotope values.

this technique. Similar to P. grandis, the nega-
tive d13C values in T. proterum suggest the use
of more closed environments whereas the pos-
itive d18O values suggests that this species
may have obtained most of its water through
its diet (Helliker and Ehleringer 2000; Yakir
and Sternberg 2000; Ometto et al. 2005).
Interestingly, more positive carbon isotope
values were observed in Pediomeryx. (Y.) ham-
iltoni, which is similar to the LBB horse spe-
cies. On the basis of its morphology P. (Y.) ham-
iltoni has been suggested to occupy a forested
environment (Webb et al. 1981). If this assign-
ment is correct, the more positive d13C values
may indicate a unique resource such as fruits
or leaves from the canopy. Similar results have
been observed in modern communities where
duikers have more positive d13C values com-
pared to other mammal taxa in a closed en-
vironment because they feed on canopy-de-
rived fruits rather than subcanopy foliage
(Cerling et al. 2004). Alternatively, with posi-
tive d13C values similar to the equids at LBB,
occupation of more open habitats cannot be
excluded. If this is the case, ingestion of more
fibrous forage such as grasses may have
caused the elevated carbon isotope value. In-
terestingly, the suggestion that Pediomeryx in-
habited more open habitats and incorporated
grass into its diet, based on the genus having
higher crowned teeth than other taxa within
the same family, also has been put forward
(Webb 1983). A recent study of the diets with-
in Dromomerycidae using microwear and me-
sowear analyses supports the idea that P. (Y.)
hamiltoni was an intermediate feeder and did
include grass in its diet (Semprebon et al.
2004).
Also of note in the isotopic results from LBB
are the negative d13C and d18O values of ‘‘H.’’
minima. Morphologically, this taxon is sus-
pected to have been a grazer (Webb et al.
1981), but isotopically it appears closer to
Gomphotherium and Tapirus, both ecologically
suspected to have been forest browsers. The
negative isotope values may also indicate
feeding on grass present in closed-canopy for-
ests, where it has been shown that taxa feed-
 RESOURCE PARTITIONING IN MIOCENE COMMUNITIES
ing on subcanopy foliage have negative car-
bon isotope values in tooth enamel (van der
Merwe and Medina 1991; Cerling et al. 2004)
due to CO2 recycling.
Black Hawk Ranch. It has been suggested
that the flora at BHR are typical of two habitat
types, floodplain and woodland-chaparral
(Condit 1938; Axelrod 1944). Isotopically, we
might expect that floodplain plants and mam-
mal inhabitants to show more negative carbon
isotope values than the arid-adapted plants
and animals of the chaparral. However, with
no significant differences in d13C values
among the eight sampled taxa at BHR, it is dif-
ficult to assess whether these two habitats and
the various resources in these habitats were
being partitioned. All taxa appear to be con-
strained around 211.8‰, the mean d13C value
for the locality, and the overall range in indi-
vidual d13C values is only 3.5‰. The limited
range around the locality mean d13C may im-
ply some combination of the following possi-
bilities: (1) BHR consisted of a relatively ho-
mogeneous flora, (2) taxa did not partition re-
sources because resources were abundant
(Gordon and Illius 1989), or (3) taxa parti-
tioned resources not by selecting different
plants, but by utilizing different parts of the
same plant or occupying the landscape at dif-
ferent times (Schoener 1974).
At BHR, there is much more spread in the
oxygen isotope values as compared to the car-
bon isotope values. The oreodont, Merychyus
major, has the most positive mean d18O value
at BHR, which may imply the use of more arid
habitats (woodland-chaparral). With the most
negative mean d18O values, the horse Hippa-
rion forcei may have been concentrated in less
arid habitats (floodplain). These differences
may also be due to water source, with the
more positive values indicating water intake
predominantly from leaf water and the nega-
tive oxygen isotope values indicating inges-
tion of local meteoric waters (Helliker and Eh-
leringer 2000; Yakir and Sternberg 2000;
Ometto et al. 2005). However, the variation ob-
served among taxa in d18O values may also re-
sult from metabolic variation among the taxa
(Kohn 1996; Kohn et al. 1996; Koch 1998). Giv-
en this uncertainty regarding the source of
variation in d18O values, we do not use these
201
oxygen isotope values as a conclusive indica-
tor of paleohabitat preference within these
taxa.
With regard to the initial aims stated in the
introduction, the isotopic data of mammalian
tooth enamel from taxa within C3-dominated
environments show that identification of re-
source partitioning is possible, especially at
the Love Bone Bed. Interpretation of the re-
sults becomes more problematic, because re-
source partitioning in modern C3-dominated
environments is incompletely understood. Re-
search is underway to sort out if the isotopic
differences observed among taxa are due to
resource use differences or to other factors
such as season of tooth growth (Brooks et al.
1997; Heaton 1999; Bocherens 2003; Richards
and Hedges 2003; Drucker and Bocherens
2004; Cerling et al. 2004; MacFadden and Hig-
gins 2004). For many of the taxa, the isotope
values corroborate what is currently under-
stood about their autecology, especially at LBB
(Table 4). In general, horses have more posi-
tive d13C values, implying use of more open
habitats, whereas antilocaprids, camelids, and
proboscideans have more negative values, im-
plying residence in more closed environ-
ments. However, for a few taxa, such as Pe-
diomeryx (Y.) hamiltoni and ‘‘Hemiauchenia’’
minima, the isotopic data provided unexpect-
ed results, given previously suggested ecolo-
gy. From what is currently understood about
isotopic patterns in modern C3-dominated en-
vironments (e.g., positive d13C and d18O values
in open habitats), it is possible to suggest rea-
sonable alternatives for these unexpected re-
sults, such as canopy-derived fruit consump-
tion. Although phylogenetically similar taxa
do appear to also be isotopically similar, data
were not sufficient to conclude whether the re-
source use of some taxa was, relative to other
taxa, constant over time; the data at hand do
suggest, however, that the relative values
among taxa will differ from locality to locali-
ty. The carbon isotope values at BHR were not
significantly different among taxa, whereas
clear differences among taxa were evident at
LLB. One distinguishing aspect between the
two localities is the disparity in ungulate di-
versity, which may explain why significant
differences are observed at LBB and not at
202 ROBERT S. FERANEC AND BRUCE J. MAC FADDEN
BHR. Only nine ungulate taxa are known
from the BHR locality (Prothero and Tedford
2000). This contrasts LBB, which currently has
18 known ungulates. Further, the LBB taxa are
suggested to have occurred in at least three
different types of habitat (Webb et al. 1981),
whereas only two habitat types are described
for BHR (Condit 1938; Axelrod 1944), and no
data exist to distinguish if taxa utilized one
particular habitat or both. Recognition of re-
source partitioning at LBB may be due to its
higher taxonomic and habitat diversity rela-
tive to BHR. Distinguishing whether the high-
er ungulate diversity at LBB necessitates re-
source partitioning, assuming a diversity
threshold was crossed after eight taxa assem-
bled within the LBB ecosystem, is not cur-
rently possible with isotopic techniques. It is
interesting to note that there is only one com-
bination of eight taxa out of 6435 total com-
binations of the 15 sampled taxa from LBB in
which no significant differences in d13C values
would be observed. Further isotopic analyses
of taxa from localities prior to 7 Ma and the
rapid increase in C4 ecosystems will aid in de-
termining how important ecosystem diversity
is to resource partitioning in ancient ecosys-
tems.
Conclusions
At both localities, horses generally have
more positive d13C values, implying use of
more open habitats, whereas antilocaprids,
camelids, and proboscideans have more neg-
ative values, implying residence in more
closed environments. The data from the Love
Bone Bed locality reveal that it is possible to
discriminate resource use differences among
taxa through the analysis of stable isotope val-
ues in tooth enamel prior to the global C4
plant expansion. The positive d13C values of P.
(Y.) hamiltoni at LBB in this study as well as
microwear and mesowear data from another
study suggest the incorporation of more fi-
brous plants, probably grass, in more open
habitats. The negative d13C values of ‘‘H.’’ min-
ima at LBB imply foraging of closed-canopy
grasses or browse rather than feeding in more
open country. The lack of significant differ-
ences among taxa at Black Hawk Ranch indi-
cates the possibility of a relatively homoge-
neous flora, the presence of abundant resourc-
es permitting niche overlap, or partitioning of
similar foods or habitat based on using the
plants at different times or using different
parts of the same plant. The results from this
study point to possible applications to discern
resource partitioning for the millions of years
before the rapid increase in C4 ecosystems that
occurred during the late Miocene.
Acknowledgments
We would like to thank A. Barnosky, P.
Koch, E. Hadly, W. Clemens, R. Amundson, R.
Hulbert, T. Dawson, M. Carrasco, J. Blois, J.
Bruzgul, Y. Chan, E. Davis, K. Feranec, D. Fox,
S. Hopkins, M. Kohn, B. Kraatz, K. O’Keefe, A.
Shabel, and P. Spaeth for discussion and com-
ments on previous versions of this manu-
script. Funding for this project was provided
by the Society of Vertebrate Paleontology’s
Predoctoral Grant, the Department of Integra-
tive Biology at the University of California,
Berkeley, the University of California Museum
of Paleontology, and a National Science Foun-
dation Doctoral Dissertation Improvement
Grant to R.S.F. Samples were provided by the
University of California Museum of Paleon-
tology and the Florida Museum of Natural
History. R. Hulbert (FLMNH) and P. Holroyd
(UCMP) provided invaluable assistance with-
in the museum collections. This is the Univer-
sity of California Museum of Paleontology
Contribution to Paleontology No. 1905.
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