Oecologia (2002) 133:14–18
DOI 10.1007/s00442-002-0977-6
In Phillips and Koch (2002), we presented an isotopic
mixing model for use in cases where elemental concen-
tration differs greatly among potential contributors to the
mixture. Our primary goal was to present the mathemat-
ics of this system for two isotopes and three sources, not-
ing that it was generalizable to n isotopes and n+1 sourc-
es. To illustrate the magnitude of the differences that
might result if elemental concentrations differed greatly
among possible sources, we performed model sensitivity
tests and examined examples involving mixing of differ-
ent food sources in the diets of captive mink and wild
bears. The response by Robbins et al. (2002) deals large-
ly with our bear example and it makes five points we
will address.
First, Robbins et al. (2002) argue that our example in-
volving bears from the Kenai Pennisula, Alaska, which
used data from their previously published work (Hilder-
brand et al. 1996; Jacoby et al. 1999), is unrealistic in
several ways, and therefore, that our results should not
be used to make management decisions. We agree com-
pletely. We did not attempt to present a rigorous re-anal-
ysis of isotopic constraints on Kenai bear diets. Our only
goal with the illustrative examples was to alert workers
to an important complication that could crop up in die-
tary (and other) isotope analyses, and to offer a tool for
addressing this complication in a quantitative manner.
Second, in addition to their concerns about our re-
sults for the coastal Kenai bears, Robbins et al. (2002)
take issue with the single sentence in our paper on in-
land bear populations. We suggested that standard linear
mixing “may be overestimating the amount of meat in
bear diets simply because bear tissue δ
15
N values will
strongly resemble those of terrestrial meat once this N-
rich source makes up more than 10 or 20% of assimilat-
ed biomass”. Robbins et al. (2002) find this suggestion
“baseless”, citing data from a subset of the inland popu-
lations with isotopic data suggesting nearly 100% plant
diets. Yet these populations have no access to salmon
and little access to large ungulates and, therefore, it is
not especially surprising that they are essentially herbiv-
orous. The existence of entirely herbivorous bear popu-
lations does not address the point we were raising.
There are populations of brown (Ursus arctos) and
black (U. americanus) bears that have access to terres-
trial meat and apparently eat it. For the Greater Yellow-
stone ecosystem, Jacoby et al. (1999) used standard lin-
ear mixing and estimated that brown bears have a mean
of 58% terrestrial meat in their diets, whereas black
bears average 48% (all data from their Table 1). Like-
wise, brown bears from the southwestern United States
have 88% terrestrial meat in their diets; black bears av-
erage 39%. Brown bears from the Blackfeet and Flat-
head Indian Reservations in Montana have estimated
meat intakes of 69%. Clearly it is only for these popula-
tions, where there is a mixture of meat and plants in the
diet, that our comments about biases in isotope mixing
are relevant. Indeed, Jacoby et al. (1999) were surprised
by the high dietary meat estimate for southwestern Unit-
ed States bears, and devoted two paragraphs to attempts
to explain the result. We stand by our suggestion that
concentration differences among leafy plants, nuts, and
terrestrial meat should be investigated as a potential
contributor to high dietary meat estimates.
Third, Robbins et al. (2002) present a strong case
demonstrating that Kenai bears consume leafy plants
rather than fruit, and argue that our model might yield
different results with a more realistic leafy plant diet.
They note that leafy plants tend to have higher N con-
centrations ([N]) and lower C:N ratios than fruit. And
because plant protein digestion is efficient, whereas di-
gestion of structural carbohydrates in leafy plants is not,
the C:N ratio of assimilated leafy plant biomass may be
P.L. Koch (
✉
)
Department of Earth Sciences, University of California,
1156 High St., Santa Cruz, CA 95064, USA
e-mail: pkoch@es.ucsc.edu
Fax: +1-831-4593074
D.L. Phillips
U.S. Environmental Protection Agency,
National Health and Environmental Effects Research Laboratory,
200 SW 35th St., Corvallis, OR 97333, USA
STABLE ISOTOPE ECOLOGY
Paul L. Koch · Donald L. Phillips
Incorporating concentration dependence in stable isotope mixing
models: a reply to Robbins, Hilderbrand and Farley (2002)
Received: 4 January 2002 / Accepted: 29 April 2002 / Published online: 30 July 2002
© Springer-Verlag 2002