Palaeogeography, Palaeoclimatology, Palaeoecology 329-330 (2012) 118–123

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Palaeogeography, Palaeoclimatology, Palaeoecology

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The carnivoran fauna of Rancho La Brea: Average or aberrant?

Brianna K. McHorse a, b, John D. Orcutt c, d, Edward B. Davis c, d,⁎
a
Department of Biology, 1210 University of Oregon, Eugene, OR 97403, USA
b
Robert D. Clark Honors College, 1293 University of Oregon, Eugene, OR 97403, USA
c
Department of Geological Sciences, 1272 University of Oregon, Eugene, OR 97403, USA
d
University of Oregon Museum of Natural and Cultural History, 1680 E. 15th Ave. Eugene, OR 97403, USA

a r t i c l e i n f o a b s t r a c t

Article history: The late Pleistocene asphalt seeps of Rancho La Brea are well-known for their impressive assemblage of carnivor-
Received 15 December 2011 ans, which make up the vast majority of the preserved fauna. Of particular interest is the large number of dire
Received in revised form 14 February 2012 wolf and sabertooth cat specimens. Carcass domination, the hypothesis that predators engaged in intense com-
Accepted 15 February 2012
petition for trapped prey, may explain the mechanism of this predator trap. Large and social animals would have
Available online 22 February 2012
fared best during competition over carcasses, so the preponderance of Canis dirus and Smilodon fatalis has been
Keywords:
seen as evidence of their sociality. However, no studies have quantitatively determined whether the relative
Rancho La Brea carnivoran species abundances in Rancho La Brea differ significantly from those in California or North America
Smilodon at large. We compare numbers of identified specimens (NISP) from the Rancho La Brea fauna to regional and
Carnivore sociality continental faunal data compiled from the FAUNMAP II database to test this hypothesis. Our results confirm
Preservation that the carnivoran fauna in Rancho La Brea is unique, with preservation patterns generally supporting the
Carcass domination carcass domination hypothesis as well as the sociality of S. fatalis.
Canis dirus © 2012 Elsevier B.V. All rights reserved.

1. Introduction biology playback experiments, where wildlife biologists play recorded

The Rancho La Brea (RLB) tar seeps are famous as a predator trap
deposit for their spectacular carnivoran preservation, particularly of
thousands of dire wolves (Canis dirus) and sabertooth cats (Smilodon
fatalis; Stock and Harris, 1992). Other members of the carnivoran
fauna include short-faced bears (Arctodus simus), the American lion
(Panthera atrox), the scimitar cat (Homotherium serum), and many
small carnivorans such as mephitids and mustelids (Carbone et al.,
2009).
Most studies suggest that the abundance of carnivorans at RLB stems
from their attraction to trapped, dying herbivores (Merriam, 1911;
Shaw and Quinn, 1986; and Stock and Harris, 1992). Spencer et al.
(2003) found evidence that carnivorans were using trapped animals
as a food resource, as their investigation of the taphonomy of the site
showed considerable ravaging of the assemblage by carnivores.
Carbone et al. (2009) suggested a carcass domination scenario to
explain the relative abundances of carnivoran taxa: large and social
predators would be most successful at defending carcasses of trapped
prey from other predators and, by extension, were most likely to be-
come trapped and preserved themselves. In contrast, solitary animals,
especially those with a small mass, would tend to avoid competing for
trapped carcasses. Carbone et al. (2009) compared RLB carnivoran
abundances directly to carnivoran abundances from modern wildlife
⁎ Corresponding author at: Department of Geological Sciences, 1272 University of
Oregon, Eugene, OR 97403, USA. Tel.: + 1 541 346 3461.
E-mail address: edavis@uoregon.edu (E.B. Davis).
sounds of dying herbivores to attract curious predators (Mills et al.,
2001). The proportions of carnivorans at RLB were not significantly
different from the proportions of ecological analogs in modern African
savannah playback studies. Comparing African lions (Panthera leo) to
sabertooth cats, hyenas to dire wolves, and jackals to coyotes led to
the conclusion that S. fatalis was likely social because of parallels be-
tween its relative abundance and that of the social P. leo in playback
experiments.
Kiffner (2009) criticized the conclusions of Carbone et al. (2009),
noting that the ecology of RLB and its carnivoran fauna are not neces-
sarily comparable to the ecology and carnivoran fauna of the modern
African savannah. This criticism is similar to the debate that has sur-
rounded “Pleistocene Rewilding,” the idea that modern African mega-
fauna should be introduced to North America to restore a Pleistocene
ecosystem (Donlan et al., 2005). Pleistocene Rewilding has been pro-
moted on the merit of ecological analogs filling empty ecosystem
roles (Donlan et al., 2006), but reservations have been raised about
the validity of substituting these analogs (Rubenstein et al., 2006).
One constructive result of this controversy is a push for exploration
of the limits of ecological analogy vs. true ecosystem function (Caro,
2007).
In this vein, instead of using modern African distributions of ecolog-
ical analogs to generate a relative abundance null hypothesis as in the
study of Carbone et al. (2009), we use the FAUNMAP II database
(Graham and Lundelius, 2010) to collect relative abundance data for
southern California (SC) and North America (NA) as a whole. We fol-
lowed Carbone et al. (2009) in including only mammals of the order

0031-0182/$ – see front matter © 2012 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2012.02.022
 B.K. McHorse et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 329-330 (2012) 118–123
Carnivora. We then quantitatively compare the RLB fauna to SC and NA,
testing the hypothesis that preservation in RLB of certain species is sta-
tistically distinct from a random sample of the surrounding geographic
region. If the carcass domination model of Carbone et al. (2009) is ap-
propriate, RLB would have a poor fit with SC/NA assemblages: social
carnivores would be over-represented and solitary species would be
under-represented. If the alternative hypothesis of Kiffner (2009) is cor-
rect and the RLB assemblage depends more on the physical properties
of the asphalt deposits than on animal behavior, we would expect
over-representation to depend more on body mass than on sociality.
2. Materials and methods
2.1. Abundance data
Carnivoran abundances in the Rancho La Brea fauna, given as mini-
mum numbers of individuals (MNI), were taken from Carbone et al.
(2009; n = 3324). We used FAUNMAP II, a continent-wide database of
published faunal data from the Pliocene (5 Ma) through the recent, to
establish carnivoran abundances in SC (n = 113) and NA as a whole
(n= 2213; Graham and Lundelius, 2010). Our FAUNMAP II search, per-
formed through the NEOMAP portal (http://www.ucmp.berkeley.edu/
neomap/search.html), was restricted to the Rancholabrean land mam-
mal age (0.24 Ma–0.011 Ma). We used the EstimateS Web Service de-
veloped for MIOMAP (http://www.ucmp.berkeley.edu/miomap/use/
index.html#InteractiveMapping) to generate tables with numbers of
identified specimens (NISP) for every locality (Appendix Tables 1–3).
NISP provided slightly larger datasets for SC and NA than MNI, which
was useful in light of the considerably greater sample size in RLB. This
is a conservative use of the abundance metrics: because MNI is a more
conservative estimate of abundance, if the difference between MNI
and NISP affects our results it will decrease the signal of overrepresen-
tation in RLB. Additionally, because we are working with relative abun-
dance rather than absolute abundance, MNI and NISP should converge
on the same values with sample sizes like the ones in this study.
We restricted the geographic search area for SC by mapping the
NA localities from FAUNMAP II using the NEOMAP implementation
of BerkeleyMapper (http://www.ucmp.berkeley.edu/neomap/use.
html#InteractiveMapping) and, using the polygon tool, selected a
subset of California south of modern San Francisco and north of the
Mexican border. Our SC sample is centered on the mammalian biodi-
versity hotspot present around southern California today (Davis et al.,
2008), and is intended to capture sites that contain many of the same
species found in RLB. In an effort to include localities of the same gen-
eral ecological type, we ignored state boundaries and included adja-
cent areas of Nevada and Arizona (Appendix Table 4). Finally, we
removed all RLB localities from the NA and SC datasets. Following
Carbone et al. (2009), we excluded all non-carnivoran taxa from the
analysis. We carried out all calculations in MS Excel 2007 except the
inverse chi-square, for which we used JMP version 9 (SAS Institute
Inc, 2010).
2.2. Statistical methods
2.2.1. Confidence limits for relative abundances
After calculating the relative abundances of each taxon in the
three data sets (MNI or NISP of the species divided by the total sample
count), we applied Goodman's (1965) simultaneous confidence-
interval calculations to establish 95% confidence intervals (Calede et
al., 2011; Eq. (1)). Using Goodman's method, for large N the degree
of freedom is always 1, which increases the rate of Type II error (i.e.,
is more likely to accept the null hypothesis).
ð1−pi Þ 1=2
 
c:i:ðpi Þ ¼ B  pi  : ð1Þ
N 7.3%) and NA (7.1 ± 1.8% and 2.4 ± 1.1%; Fig. 2, Table 2).
119
In this equation, pi is the relative abundance, c.i.(pi) is the confidence
interval around the relative abundance in percentage points, N is the
total sample size, and B is equal to the inverse chi-square of the degrees
of freedom and the confidence limit (here α = 0.05) divided by number
of categories (in this case, the number of taxa present in each assem-
blage). Dividing α by the number of taxa is necessary because the
uncertainty is spread across all parts of the abundance distribution.
2.2.2. Jackknife
Because of the strong effect of the two most common taxa (C. dirus
and S. fatalis) on the overall relative abundance distribution in RLB,
we applied a jackknife to the dataset, removing those two species
before repeating the relative abundance analysis. Comparing the rela-
tive abundances of the other members of the carnivore fauna to our
null distributions without jackknifing would produce misleading re-
sults. We hypothesize, as have many before us (Stock and Harris,
1992; Spencer et al., 2003; and Carbone et al., 2009), that these two
taxa are over-represented in the sample. We are able to test this
hypothesis not only by comparing the relative abundances of C. dirus
and S. fatalis directly between RLB, SC, and NA, but also by comparing
the relative abundances of the other carnivorans in RLB to SC and NA
in the absence of C. dirus and S. fatalis. We predict that the relative abun-
dances in our jackknifed dataset will not be significantly different
between RLB, SC, and NA if the two supposedly social species are the
only species oversampled in the fauna. Additionally, if the playback
analogy of Carbone et al. (2009) is correct, we should expect to see
significantly lower representation of species that would actively avoid
the competition surrounding a dying prey animal, such as cheetah-
like felids (Miracinonyx species) and all small-bodied carnivorans (e.g.,
mephitids, mustelids, and procyonids).
Alternatively, if the physics of the asphalt deposit is the driving
factor in RLB preservation (Kiffner, 2009), we would expect to see sim-
ilar overabundance in representation of all large carnivorans, so our
jackknifed RLB dataset would contain additional large-bodied animals
that are significantly higher than SC and NA values, and all small-
bodied animals (presumably below some lower entrapment threshold)
would be under-represented.
3. Results
3.1. Relative abundance
C. dirus and S. fatalis have relative abundances of 51.2 ± 2.6% and
33.3 ± 2.4% in RLB, respectively, compared to 8.8 ± 8.0% and 6.2 ±
6.8% in Southern California (Fig. 1, Table 1). In North America as a
whole they comprise 17.1 ± 2.6% and 1.8 ± 0.9% of the carnivoran
population. For these two taxa, the differences between RLB and the
other two regions are significant. Canis latrans represents 7.7 ± 1.3%
of the RLB carnivoran fauna, making it underrepresented when com-
pared to SC (23 ± 11.9%) and approximately even with NA (5.8 ±1.6%;
Fig. 1). P. atrox is present at similar levels in RLB and NA at 2.6 ± 0.8
and 1.9 ± 1.0 of the fauna; the calculated value for P. atrox in SC is
5.3% but the uncertainty is ± 6.3%.
All remaining species have lower relative abundance values in RLB
than in both SC and NA (Table 1). The small sample size in SC contrib-
utes to considerable overlap in uncertainty between RLB and SC; con-
sequently, there are no significant differences between the relative
abundances of these remaining taxa for RLB and SC.
3.2. Jackknife
Applying a jackknife to remove S. fatalis and C. dirus from the anal-
ysis reveals significant overrepresentation in RLB of C. latrans (49.6 ±
6.5%) and P. atrox (49.6 ± 6.5%) relative to SC (27.1 ± 13.4% and 6.3 ±
120 B.K. McHorse et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 329-330 (2012) 118–123

60%

50%

Relative Abundance
40%

30%

20%

10%

0%

Fig. 1. Relative abundance with 95% confidence limits for all carnivoran species present at RLB. RLB abundances are on left, SC in middle, and NA on right. Taxa are presented in rank-
order by RLB abundance.

Clear representation patterns emerge between RLB and NA in the
absence of C. dirus and S. fatalis. A. simus, Mephitis mephitis, Canis
lupus, Urocyon cinereoargenteus, Puma concolor, Taxidea taxus, and
H. serum all show similar relative abundances in RLB and NA
(Table 2). Mustela frenata, Lynx rufus, Spilogale putorius, Panthera onca,
Ursus americanus, and Ursus arctos are all significantly less common in
RLB than in NA.
Large uncertainty values for SC, a consequence of the small sample
size, mask any potentially meaningful differences between RLB and
SC for all species besides the two most abundant in the jackknife,
C. latrans and P. atrox. This lack of significance is similar to the result
of our analysis of the non-jackknifed dataset.
4. Discussion
In RLB, S. fatalis and C. dirus each represent a significantly larger
proportion of the carnivoran fauna than in either SC or NA. Many
other species, including ursids and small solitary animals like L. rufus,
are significantly underrepresented in RLB compared to the surrounding
regions. This aberrant preservation allows us to reject the null hypoth-
esis that the RLB carnivoran assemblage is simply a random subsample
of the general area.
The overrepresentation of the large, social C. dirus at RLB lends
support to the carcass domination scenario. The sociality of C. dirus
is well-constrained phylogenetically because most modern canids,
especially large ones, are social (Wang and Tedford, 2010). When
the jackknife is applied, C. latrans and P. atrox are also significantly
overrepresented in RLB compared to the controls—despite consider-
able uncertainty values for SC. C. latrans is classified as small by
Carbone et al. (2009), but modern coyotes are frequently social,
usually maintaining pair bonds or small packs that can bring down
larger prey. They also occasionally form much larger groups around
carrion (Nowak and Paradiso, 1999). Social behavior, especially forming
aggregations around carcasses, would again be an advantage if preda-
tors were competing for trapped prey in the asphalt seeps. P. atrox has
generally been considered social on the basis of its close relation to
P. leo and its pronounced sexual dimorphism (Carbone et al., 2009;
Wheeler and Jefferson, 2009; and Meachen-Samuels and Binder,
2010). Some research has suggested that P. atrox is most closely related
to, and possibly behaviorally similar to, P. onca (Christiansen and Harris,
2009); however, modern DNA evidence has conclusively demonstrated
that the closest living relative of P. atrox is P. leo (Barnett et al., 2009).
The significant difference between P. atrox and P. onca representation
at RLB further supports behavioral dissimilarity. While the sociality of
S. fatalis is still under debate (McCall et al., 2003 and Kiffner, 2009), it
is significantly overrepresented in the RLB population, falling out with
other highly social species. Our results support the conclusions of
Carbone et al. (2009) that S. fatalis was likely social.
Though C. lupus is large and highly social, its representation in RLB
is not comparable to the other large, social species. After jackknifing,

Table 1
Relative abundance values and 95% confidence intervals. Column on left notes how relative abundance in RLB compares to NA and SC.

Taxon RLB% ±95% c.i. SC% ± 95% c.i. NA% ±95% c.i.

Overrepresented Canis dirus 51.20 2.58 8.85 8.04 17.13 2.63

Smilodon fatalis 33.33 2.43 6.19 6.82 1.76 0.92
Underrepresented Canis latrans 7.67 1.37 23.01 11.91 5.78 1.63
Panthera atrox 2.59 0.82 5.31 6.34 1.94 0.96
Arctodus simus 0.99 0.51 2.65 4.55 4.61 1.46
Mephitis mephitis 0.96 0.50 1.77 3.73 4.61 1.46
Mustela frenata 0.57 0.39 5.31 6.34 6.46 1.71
Canis lupus 0.54 0.38 2.65 4.55 2.67 1.12
Urocyon cinereoargenteus 0.48 0.36 7.08 7.26 3.16 1.22
Puma concolor 0.45 0.35 4.42 5.82 1.27 0.78
Taxidea taxus 0.39 0.32 5.31 6.34 2.71 1.13
Lynx rufus 0.27 0.27 6.19 6.82 4.29 1.41
Spilogale putorius 0.18 0.22 3.54 5.23 4.16 1.39
Panthera onca 0.15 0.20 0.00 0.00 2.62 1.11
Homotherium serum 0.15 0.20 0.00 0.00 0.59 0.53
Ursus americanus 0.03 0.09 3.54 5.23 8.95 1.99
Ursus arctos 0.03 0.09 0.88 2.65 1.40 0.82
 B.K. McHorse et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 329-330 (2012) 118–123 121

60%

50%

Relative Abundance
40%

30%

20%

10%

0%

Fig. 2. Jackknife analysis: relative abundances with uncertainty when C. dirus and S. fatalis are removed from the analysis. Order of species and bars as in Fig. 1.

the relative abundance of C. lupus is not significantly different between
RLB, SC, and NA. This proportion of gray wolves fits the null hypothesis
of random sampling, and clearly contradicts the carcass domination
hypothesis. The background proportions of C. lupus could stem from
their ecological similarity to the much larger C. dirus, which may have
locally displaced C. lupus (Leonard et al., 2007). Carbone et al. (2009)
noted the similarity of C. lupus to the African wild dog, Lycaon pictus,
which is also large, social, and relatively rare in playback experiments.
It would be necessary to conduct a more detailed study of the relative
representation of C. dirus and C. lupus in Rancholabrean faunas across
North America to establish whether these species experienced compet-
itive displacement.
Taxa without significant differences in representation between
RLB and NA after jackknifing are either large and solitary or small. If
a combination of large size and sociality contribute to the likelihood
that a species will be more relatively abundant at RLB than across
NA, it follows that species possessing one or the other trait will be
less abundant than those with both. The behavior of H. serum is not
certain; some have suggested that Homotherium was likely social
because it hunted large prey (Antón et al., 2005) but others note
that the massive cat is so rare in the fossil record it is likely to have
been solitary (Carbone et al., 2009). The fact that such a rare predator
appears in the RLB assemblage at all (RLB n = 5, SC n = 0, NA n = 13)
may support sociality; it is certainly more-represented at RLB than in
the surrounding area, and RLB contains nearly half as many speci-
mens as the whole of our North American sample.
Following the same pattern, M. frenata, L. rufus, and S. putorius, all
small and solitary, make up significantly less of the carnivoran fauna
in RLB than NA. It is likely that, in the face of competition for carcasses,
these smaller and nonsocial animals would avoid the site entirely.
However, also underrepresented in RLB are P. onca, U. americanus, and
U. arctos, all large and solitary. The ursids are most striking for
the large difference between their relative abundances in RLB (0.2 ±
0.6% for both) and NA (11.0 ± 2.2% for U. americanus and 1.7 ± 0.9%
for U. arctos). Diet is likely a major reason for this discrepancy; unlike
most large taxa in the RLB carnivoran fauna, these ursids tend toward
omnivory, and today upwards of 80% of their diet can consist of plant
matter and invertebrates (Mattson, 1998). They may have avoided the
asphalt seeps, unwilling to engage in such competition for the carcasses
when they could satisfy their nutritional requirements elsewhere.
The possibility remains that factors other than carcass competition
played a role in RLB's atypical faunal representation. Kiffner (2009)
suggested that size may have had an effect on the likelihood of an
animal becoming trapped in the tar, with larger animals more likely
to become fatally stuck. The dearth of large herbivores at RLB suggests
that size was not the main factor (Van Valkenburgh et al., 2009), but
testing the effects of body mass in different viscosities of fluid could
help suggest whether size may have contributed to the aberrant

Table 2
Relative abundance values and 95% confidence intervals after jackknifing. Column on left notes how relative abundance in RLB compares to NA and SC.

Taxon RLB% ±95% c.i. SC% ±95% c.i. NA% ±95% c.i.

Overrepresented Canis latrans 49.61 6.47 27.08 13.49 7.13 1.78

Panthera atrox 16.73 4.83 6.25 7.35 2.40 1.06
Equally-represented Arctodus simus 6.42 3.17 3.13 5.28 5.68 1.60
Mephitis mephitis 6.23 3.13 2.08 4.33 5.68 1.60
Mustela frenata 3.70 2.44 6.25 7.35 7.97 1.88
Canis lupus 3.50 2.38 3.13 5.28 3.29 1.24
Urocyon cinereoargenteus 3.11 2.25 8.33 8.39 3.90 1.34
Puma concolor 2.92 2.18 5.21 6.74 1.56 0.86
Taxidea taxus 2.53 2.03 6.25 7.35 3.34 1.25
Homotherium serum 1.75 1.70 7.29 7.89 5.29 1.55
Underrepresented Lynx rufus 1.17 1.39 4.17 6.07 5.13 1.53
Spilogale putorius 0.97 1.27 0.00 0.00 3.23 1.23
Panthera onca 0.97 1.27 0.00 0.00 0.72 0.59
Ursus americanus 0.19 0.57 4.17 6.07 11.03 2.17
Ursus arctos 0.19 0.57 1.04 3.08 1.73 0.90
122 B.K. McHorse et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 329-330 (2012) 118–123
composition of the RLB assemblage. Our data support both size- and
sociality-influenced entrapment rates rather than size alone. The
most poorly represented carnivoran taxa in RLB are the massive but
solitary ursids. The significant under-representation of ursids sug-
gests grounds for rejection of the model for RLB preservation that
depends solely on the physical properties of the asphalt. If physical
entrapment alone, with no carcass competition, were the dominant
process building the RLB sample, we would expect to see all large-
bodied species better represented than the null expectation, as their
weight would have caused any interactions with the tar to be more
likely to result in capture and preservation. Instead, we see support
for the carcass domination hypothesis, with large, social carnivorans
drawn to the dying herbivores, but large solitary species driven away.
The small sample size in SC after removal of RLB localities proved
problematic, creating uncertainty that was in some cases greater
than the relative abundance values themselves. This high degree of
uncertainty precluded direct comparisons with RLB for all but the
four best-represented taxa. More extensive sampling would resolve
this problem, producing numbers of specimens that would allow a
more robust comparison to surrounding areas rather than NA as a
whole. Following Moore et al. (2007), a sample greater than 534 indi-
viduals would provide confidence limits within 5% of observed values.
Despite this limitation, the values for C. dirus, S. fatalis, C. latrans, and
P. atrox were significantly different between RLB and SC, providing
quantitative confirmation of atypical faunal preservation in RLB relative
to the surrounding area.
While concerns about small sample size do not apply to the
Rancholabrean fauna of North America, there are potential biases that
may explain, at least in part, the differences in relative abundance
between RLB and the continent as a whole. Chief among these is the
preponderance of cave sites in the North American record (Graham
and Lundelius, 2010). Because of the extraordinary preservational
conditions present in most caves, this preponderance might reasonably
be expected to skew the relative abundances of certain taxa at a conti-
nental scale. This might especially be the case for carnivores, as many
caves are thought to represent predator traps as well (Wang and
Martin, 1993). However, Muñoz-Durán and Van Valkenburgh (2006)
showed that cave deposits actually represent a more accurate snapshot
of Rancholabrean carnivorans diversity than do surface deposits, where
small-bodied taxa in particular are underrepresented. Thus the prepon-
derance of cave sites should actually reduce biases in carnivore relative
abundance at a continental scale. This further underscores the point
that RLB is a taphonomically unique and biased locality that is not a
good ecological analog for the standing paleocommunity.
5. Conclusions
The preservation of carnivorans at Rancho La Brea (RLB) is aberrant
when compared to the surrounding geographic region (Southern
California, SC) and to North America (NA) as a whole. Comparison
between RLB and SC was limited by the small sample size of SC, except
in the four most-represented taxa. Smilodon fatalis, Canis dirus, Canis
latrans, and Panthera atrox have significantly greater relative abun-
dances in RLB than predicted by the null hypothesis. Three of the
over-represented species are large and probably social, and the other,
C. latrans, is small and social. Large, solitary taxa and other small, social
carnivoran species have relative abundances in RLB that are, for the
most part, not significantly different from those in the surrounding
areas. The least-represented species at RLB are mainly small and soli-
tary, with the notable exception of ursids (bears), which are large and
solitary. The ursid underrepresentation is unexpected under hypothe-
ses of purely physical means of sampling by the tar seeps: if the physical
properties of the tar controlled sampling, we would expect ursids to be
as over-represented as other carnivorans in their size class. We suggest
that ursid under-representation may instead be related to their
omnivorous diet, supporting the carcass domination hypothesis of
Carbone et al. (2009).
Together, our results support the carcass domination hypothesis,
which suggests that large, social carnivorans are over-represented in
the RLB fauna because they could successfully compete for access to
trapped prey, and, as a consequence, were more likely to be trapped
themselves. We reject the null hypothesis that the carnivoran fauna
of Rancho La Brea is simply a random subsample of the faunas of
surrounding geographic areas.
Supplementary materials related to this article can be found
online at doi:10.1016/j.palaeo.2012.02.022.
Acknowledgments
We thank S. Hopkins for constructive discussion and supporting
facilities. We also thank the members of the Hopkins lab, J. Calede, W.
Kehl, A. Gusey, A. Atwater, W. McLaughlin, K. Stilson, and D. Levering,
for their feedback. We would also like to thank two anonymous
reviewers whose critical feedback substantially improved an earlier
version of this manuscript. This work was completed while B.M. was
supported by the Singer Foundation, University of Oregon, UO Robert
D. Clark Honors College, and UO Department of Biology. The Society of
Vertebrate Paleontology provided J.O. funding to travel to the society's
annual meeting to present and discuss the preliminary results of this
research.
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