Functional Ecology 2015, 29, 34
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FE SPOTLIGHT
Nutritional geometry provides new insights into the
interaction between food quality and demography in
endangered wildlife
Jessica M. Rothman*
Department of Anthropology, Hunter College of the City University of New York, 695 Park Avenue, New York 10065,
USA
Faced with seasonally and spatially heterogeneous land-
scapes, wildlife need to make careful foraging decisions to
consume adequate amounts and concentrations of essential
macro- and micronutrients. These decisions vary as a func-
tion of ontogeny, thermoregulation and physiological
states; for example, growing juveniles and lactating females
must meet greater energy and protein requirements per
kilogram of metabolic body mass than males. Disentan-
gling the demographic constraints imposed by food quality
on endangered wildlife is particularly pressing and vital to
developing eective conservation plans that support
successful reproduction.
Wildlife must balance multiple nutrients in a manner that
enhances tness. Theoretical frameworks need to consider
that nutritional needs are dynamic and complex, and ani-
mals need to full a variety of nutritional needs simulta-
neously (Raubenheimer, Simpson & Mayntz 2009). While a
number frameworks have been developed to interpret the
nutritional goals of wildlife foraging, such as optimal for-
aging theory (Stephens & Krebs 1986) and protein maximi-
zation (White 1993), these are typically single-currency
models that assume animals will maximize the acquisition
of a particular nutrient such as protein or energy. The geo-
metric framework (GF) is a multidimensional state-depen-
dent framework that considers nutritional decisions of
animals based on interactions among nutrients and toxins
as foods are consumed (Simpson & Raubenheimer 2012).
Using Cartesian geometry, the GF is designed to decipher
nutritional interactions through a nutrient space built of
two or more axes, whereby axes represent dietary nutrients
or toxins expected to be functionally signicant. As the ani-
mal consumes a diet, its position in nutrient space changes
according to the nutrients consumed, and its responses to
multiple dietary nutrients can be explored. Newly devel-
oped right-angle mixture triangles (n-dimensional plots of
compositional data) are a useful tool of the GF to visually
explore relationships among dierent proportions of nutri-
ents and toxins within food in relation to parameters such
*Correspondence author. E-mail: jessica.rothman
@hunter.cuny.edu
2015 The Author. Functional Ecology 2015 British Ecological Society
doi: 10.1111/1365-2435.
as nutrient requirements, digestive eciencies or environ-
mental availability (Raubenheimer 2011).
In the current issue of Functional Ecology, Nie et al.
(2014) used the GF to make a major leap forward in our
understanding of wildlife nutritional ecology, specically
linking food quality to demography in free-ranging endan-
gered giant pandas that reside in China. Female pandas
only have three or four surviving ospring in their lifetime;
thus, understanding the eects of nutritional constraints
on successful reproduction and ospring survival is criti-
cal. Giant pandas are unique in their feeding ecology
because they are specialist herbivores but are in the Order
Carnivora. Like other members of Carnivora, they have
large canine teeth, and a short and simple digestive tract.
Consequently, they are inecient in digesting their food
compared to other herbivores (Dierenfeld et al. 1982).
Using GF, Nie and colleagues demonstrated that wild pan-
das shift their migrations in response to seasonal changes
in availability and nutrient quality of bamboo, their princi-
ple food item. Pandas favoured nitrogen-rich young shoots
over leaves when shoots were available in the springtime.
As the shoots matured and had reduced in concentrations
of nitrogen, leaves were preferred. While young bamboo
shoots are higher in N than leaves, shoots have a lower
ratio of Ca to P than leaves so much lower that the
authors suggest that it would not allow for adequate bone
growth. As a result, pandas may only receive an adequate
Ca to P ratio when they are eating leaves. By switching
diets seasonally, pandas synchronize reproduction with
eating N-rich shoots in the springtime, possibly allowing
for placental tissue growth, and delay implantation until
they eat bamboo leaves to allow for Ca investment in
lactation and bone growth.
Importantly, Nie et al. (2014) demonstrated that intra-
specic shifts in the nutritional composition of bamboo
shoots and leaves correspond to seasonal migrations and
feeding switches that coincide with reproductive seasonali-
ty in pandas. Pandas migrated to dierent areas of their
habitat seasonally, gaining complementary nutrients from
each, indicating that these dierent habitats are not
nutritionally interchangeable. These results illustrate the
utility of robust and explicit frameworks such as GF for
4 FE Spotlight
implementing conservation tactics. Current conservation
plans of pandas and other wildlife focus on habitat suit-
ability (Loucks et al. 2001; Zhang et al. 2011; Sawyer &
Brashares 2013); the results provided by Nei and col-
leagues suggest that knowledge of nutritional phenology
should also be incorporated to enhance wildlife tness.
This study is the rst to use GF to link the feeding
strategies of an endangered herbivore to its demography.
Previously, the GF has been used to assess the nutritional
decisions of captive animals (e.g., Behmer & Joern 2008;
Dussutour et al. 2010; Kohler, Raubenheimer & Nichol-
son 2012), and only recently has it been used to interpret
the nutritional goals of animals in wild environments
including mountain gorillas (Rothman, Raubenheimer &
Chapman 2011), grizzly bears (Coogan et al. 2014), spider
monkeys (Felton et al. 2009) and chacma baboons
(Johnson et al. 2013). In addition to using multidimen-
sional frameworks to unravel the nutritional challenges of
wildlife in relation to their demography (DeGabriel et al.
2013), a future priority of nutritional ecologists should
include expanding the chemical data base of wildlife foods
over seasonally and spatially explicit landscapes and inte-
grating these measures with estimates of food intake when-
ever possible. Understanding these nutritional challenges is
even more pressing as a result of global change and the
interactive eects of CO2, increasing temperature and
rainfall variability on both food quantity and quality.
Acknowledgements
I am grateful to Chuck Fox for inviting me to write this commentary, and I
thank Joanna Lambert and Linda DAnna for their constructive comments
on an earlier draft.
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