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Stable Isotope Techniques and Applications for Primatologists

Stable Isotope Techniques and Applications for Primatologists

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Stable Isotope Techniques and Applicationsfor Primatologists
Brooke E. Crowley
Received: 18 February 2011 /Accepted: 18 October 2011
#
Springer Science+Business Media, LLC 2012
Abstract
Stable isotope biogeochemistry is useful for quantifying the feeding ecol-ogy of modern and extinct primates. Over the past three decades, substantial advanceshave been made in our knowledge of the physiological causes of isotopic patterns aswell as effective methodology to prepare samples for isotopic analysis. Despite theseadvances, the potential of stable isotope biogeochemistry has yet to be fully exploited by primate researchers, perhaps due to the prolific and somewhat daunting nature of the isotopic literature. I here aim to present a cogent overview of stable isotopeapplications to nonhuman primate feeding ecology. I review the factors that influenceecological patterns in carbon, nitrogen, and oxygen stable isotopes. I present methodsfor collecting and preparing samples of tooth enamel and bone mineral hydroxyapatite, bonecollagen, fur and hairkeratin, blood,feces, and urine for isotope analysis. I discuss both the existing and potential applications of these isotopic patterns to primate feedingecology. Lastly,I point out some ofthepitfalls to avoidwheninterpretingand comparingisotopic results.
Keywords
δ
13
C.
δ
15
 N.
δ
18
O.Hydroxyapatite.Proteinaceoustissues.Stableisotope.Trophicdiscrimination
Introduction
Stable isotope biogeochemistry is a powerful tool for enhancing our understanding of the diets and niche partitioning of extant species as well as reconstructing the dietsand habitats of extinct species. Our current knowledge of nonhuman primate dietaryneeds derives largely from field observations, dental wear, and morphology. Al-
Int J PrimatolDOI 10.1007/s10764-012-9582-7
Electronic supplementary material
The online version of this article (doi:10.1007/s10764-012-9582-7)contains supplementary material, which is available to authorized users.B. E. Crowley (
*
)Departments of Geology and Anthropology, University of Cincinnati, Cincinnati, OH 45221, USAe-mail: brooke.crowley@uc.edu
 
though each of these methods has its merits, each is also inherently incomplete(Sponheimer 
et al.
2009). Observed and actual feeding rates may differ becausemany species are cryptic and difficult to study. Dental wear may be biased towardgritty or acidic foods and may only reflect an animal
s most recent meals (Grine1986). Morphology clearly reflects dietary adaptations, but such adaptations might bemore influenced by the consumption of fallback foods (Lambert 
et al.
2004; Marshalland Wrangham2007). As a result, actual feeding ecology may be much broader thanmorphological adaptations would suggest. Isotopic data, providing a quantitativerecord of an animal
s feeding ecology, can validate observational and morphologicaldata sets.The application of stable isotope biogeochemistry to ecological research has beenincreasing steadily over the last three decades. Awealth of papers on topics includinghabitat characterization, plant and animal physiology, and food web dynamics is nowavailable. However, this information is not always accessible to those unfamiliar withthe isotopic literature. Here I provide background information on carbon, nitrogen,and oxygen isotopes, the three stable isotope systems that are most commonly used infeeding ecology studies. I review collection and sample preparation methods for thetissues and body products most relevant to primate feeding ecology (tooth enamel and bone hydroxyapatite, bone collagen, fur and hair keratin, blood, feces and urine), and present some examples of stable isotope applications in primate foraging ecology.Stable isotope biogeochemistry has been widely applied in archaeological studies.Whereas summarizing this literature is beyond the scope of this article, I do refer tohuman-based studies when examples for nonhuman primates are lacking.A number of excellent recent reviews cover the fundamentals of isotope biogeo-chemistry in plants (Barbour 2007; Handley
et al.
1999; Heaton1999; Kohn2010; Marshall
et al.
2007) and animals (Gannes
et al.
1998; Koch2007; Kohn and Cerling 2002; Lee-Thorp
et al.
2003; Martínez del Rio
et al.
2009; Schoeninger 2010; Sponheimer 
et al.
2009). I refer readers to these sources for a more in-depthdiscussion of the material presented here. Isotopic analysis of additional tissues, e.g.,tooth dentine, and additional isotopes, e.g., hydrogen, strontium, and sulfur, may alsohave applications to primate foraging ecology (Gannes
et al.
1998; Koch2007). However, discussion of these topics is beyond the scope of this article.
Background on Stable Isotope Biogeochemistry
Stable isotope ratios in animal tissues are informative because they provide a recordof the food or water actually ingested and assimilated into an animal
s body tissues.These ratios are predicted to vary as a function of habitat, physiology, and dietary proclivity (Koch2007; Schoeninger 2010). Accordingly, stable isotopes have been used to investigate foraging ecology and niche partitioning among both living andextinct nonhuman primates (Codron
et al.
2006; Crowley
et al.
2011a ; Dammhahnand Kappeler 2010; Fourie
et al.
2008; Lee-Thorp
et al.
2003; Loudon
et al.
2007;Schoeninger 
et al.
1997).Stable isotope ratios are ideal for these purposes because the difference in mass between isotopes of an element is measurable. Isotopes with more neutrons, e.g.,
13
C,
15
 N, and
18
O, have heavier masses than isotopes with fewer neutrons, e.g.,
12
C,
14
 N,
B.E. Crowley
 
and
16
O. These mass differences result in slightly different thermodynamic andkinetic properties among isotopes of the same element (Hoefs1997; Urey1947). They also lead to partitioning, or fractionation, of lighter and heavier isotopes between different materials during physical and chemical reactions (Hoefs1997).As a result of these fractionation processes, different materials have discrete isotoperatios, reflecting the relative amounts of lighter and heavier isotopes. Becausethe absolute ratio of heavy to light isotope tends to be very small, isotopicabundance is generally referred to as parts per thousand (per mil,
) inreference to an international standard using a standardized
δ
notation, e.g.,
δ
13
C, where
¼
R
sample
=
 R
standard
À Á
À À
1
À Á
Ã
1000, and
R
is the ratio of heavy tolight isotope (
13
C/ 
12
C,
15
 N/ 
14
 N or 
18
O/ 
16
O). Positive and negative
δ
values indicatethat the sample has more or less of the heavy isotope relative to the standard,respectively (Hoefs1997).Whereas nitrogen and carbon isotopes have only one international standard,oxygen has two:
δ
15
 N is reported relative to atmospheric nitrogen (AIR);
δ
13
C isreported relative to PeeDee Belemnite (PDB), an extinct cephalopod; and
δ
18
O isreported relative to both PDB and standard mean ocean water (SMOW) (Koch2007).As a result, the scales for 
δ
values from different elements are not directlycomparable. Nevertheless, trends in
δ
13
C,
δ
15
 N, and
δ
18
O values are the same. If a material has relatively less of the heavy isotope (less
13
C,
15
 N, or 
18
O), then its
δ
value is lower. Conversely, if a substance has relatively more of the heavy isotope(more
13
C,
15
 N, or 
18
O), its
δ
value is higher. Isotope data are typically graphed as bivariate scatter plots. Carbon is conventionally placed on the
x
-axis, and nitrogen or oxygen on the
y
-axis.
Interpreting Stable Isotope Ratios in Primates
Stable isotope values in a primate
s tissues can only distinguish isotopically distinct dietary items. They usually cannot be used to determine the relative contribution of a single plant or arthropod species. Nevertheless, stable isotope values can differentiateconsumption of plant and animal matter as well as determine the relative contributionof certain plant parts, e.g., leaves versus fruits, or plant functional types, e.g., C
4
grasses, succulents, trees with N-fixing symbionts.Dietary InputsBecause isotope values in primates reflect diet, it is important to understand thefactors underlying isotopic variation in plants, which comprise the majority of most  primate diets. Carbon isotope values mainly reflect plant physiology. Plants use threemain types of photosynthetic pathways: C
3
, C
4
, and crassulacean acid metabolism(CAM). Most trees, shrubs, and grasses from regions with cool growing seasons usethe C
3
photosynthetic pathway. Grasses that grow in tropical regions or temperateregions with warm growing seasons use the C
4
photosynthetic pathway (Koch2007;Kohn and Cerling2002). The majority of leaf succulents, and many epiphytic orchidsand bromeliads, use CAM (Keeley and Rundel2003). These three plant types can bedistinguished based on their isotope values. The average
δ
13
C values for C
3
and C
4
Stable Isotope Techniques and Applications

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