American Journal of Primatology

RESEARCH ARTICLE
Fatty Acids in Mountain Gorilla Diets: Implications for Primate Nutrition and
Health
WHITNEY B. REINER1,2, CHRISTINA PETZINGER3, MICHAEL L. POWER3,4, DAVID HYEROBA5,
1,6
AND JESSICA M. ROTHMAN *
1
Hunter College of the City University of New York, New York City, New York
2
University of California Berkeley, Berkeley, California
3
Nutrition Laboratory, Smithsonian National Zoological Park, Washington, District of Columbia
4
Research Department, American College of Obstetricians and Gynecologists, Washington, District of Columbia
5
Jane Goodall Institute, Entebbe, Uganda
6
The New York Consortium in Evolutionary Primatology [NYCEP], New York City, New York

Little is known about the fatty acid composition of foods eaten by wild primates. A total of 18 staple foods
that comprise 97% of the annual dietary intake of the mountain gorilla (Gorilla beringei) were analyzed
for fatty acid concentrations. Fruits and herbaceous leaves comprise the majority of the diet, with fruits
generally having a higher mean percentage of fat (of dry matter; DM), as measured by ether extract (EE),
than herbaceous leaves (13.0%
SD 13.0% vs. 2.3
SD 0.8%). The mean daily EE intake by gorillas was
3.1% (DM). Fat provided 14% of the total dietary energy intake, and 22% of the dietary non‐protein
energy intake. Saturated fatty acids accounted for 32.4% of the total fatty acids in the diet, while
monounsaturated fatty acids accounted for 12.5% and polyunsaturated fatty acids (PUFA) accounted
for 54.6%. Both of the two essential PUFA, linoleic acid (LA, n‐6) and a‐linolenic acid (ALA, n‐3), were
found in all of the 17 staple foods containing crude fat and were among the three most predominant
fatty acids in the diet: LA (C18:2n‐6) (30.3%), palmitic acid (C16:0) (23.9%), and ALA (C18:3n‐3)
(21.2%). Herbaceous leaves had higher concentrations of ALA, while fruit was higher in LA. Fruits
provided high amounts of fatty acids, especially LA, in proportion to their intake due to the higher fat
concentrations; despite being low in fat, herbaceous leaves provided sufficient ALA due to the high
intake of these foods. As expected, we found that wild mountain gorillas consume a diet lower in EE, than
modern humans. The ratio of LA:ALA was 1.44, closer to agricultural paleolithic diets than to modern
human diets. Am. J. Primatol. © 2013 Wiley Periodicals, Inc.

Key words: nutritional ecology; essential fatty acid; linoleic; alpha‐linolenic; polyunsaturated; ape
diets

INTRODUCTION Once the essential fatty acids have been con-

Fatty acids are required components of physio-
logical mechanisms that facilitate normal neural,
retina, and brain growth, tissue development, as
well as cellular, cardiovascular, and immune
system maintenance and function [Conner, 2000;
Wainwright & Ward, 1997]. Little is known about the
fatty acid composition of the diets of wild primates,
and which foods consumed provide the largest
proportion of essential fatty acids. There are two
fatty acids commonly accepted as essential fatty
acids: linoleic acid (LA, C18:2n‐6) and a‐linolenic acid
(ALA, C18:3n‐3), which are also often referred to as
omega‐6 and omega‐3 fatty acids. The term omega
refers to the location of the first double bond from the
methyl terminal end of the fatty acid. Dietary intake
of these fatty acids is essential for primates including
humans, similar to other mammals, because we lack
the enzymes required to synthesize them [Pereira
et al., 2003].
sumed, they are converted through a series of
enzymes to their respective long‐chain derivatives.
For both the n‐6 and n‐3 fatty acids, the long‐chain
fatty acids derived from essential fatty acids have
Contract grant sponsor: Jane V. Engel Family Foundation;
contract grant sponsor: Cornell University; contract grant
sponsor: Hunter College’s Presidential Fund for Faculty
Advancement.

Correspondence to: Jessica M. Rothman, Department of
Anthropology, Hunter College of the City University of New
York, 695 Park Avenue, Room 724, New York City, NY 10065. E‐
mail: jessica.rothman@hunter.cuny.edu
Received 20 October 2012; revised 2 September 2013; revision
accepted 14 October 2013
DOI: 10.1002/ajp.22232
Published online XX Month Year in Wiley Online Library
(wileyonlinelibrary.com).

© 2013 Wiley Periodicals, Inc.

2 / Reiner et al.
been shown to play a role in immune response and the
central nervous system [Anderson & Ma, 2009]. For
example, the n‐6 fatty acid arachidonic acid (AA) is
responsible for producing eicosanoids, signaling
molecules that have a large role in immune function
and the inflammatory response. Eicosanoids are
more potent than molecules produced by the n‐3
fatty acid equivalent, eicosapentaenoic acid (EPA)
[Anderson & Ma, 2009]. Thus, diets composed of
different proportions and amounts of n‐6 and n‐3
fatty acids can alter the immune state of mammals.
Docosahexaenoic acid (DHA), another long‐chain n‐3
fatty acid, has recently been shown to also play a role
in the inflammatory status of tissues [Chapkin
et al., 2009]. DHA is also required for proper neural,
retina, and brain growth and function [Uauy &
Dangour, 2006]. Docosapentaenoic acid (DPA) is
another n‐3 fatty acid that is an important precursor
to DHA. Both DPA and DHA are critical in brain
function [Crawford, 1992; Crawford et al., 1999].
Total lipid, or fat, intake in wild primate diets has
been quantified for several primates, including
cercopithecines, chimpanzees (Pan troglodytes) [Con-
klin‐Brittain et al., 1998], saki monkeys (Pithecia
pithecia) [Norconk and Conklin‐Brittain, 2004], mon-
goose lemurs (Eulemur mongoz) [Curtis, 2004], col-
lared lemurs (Eulemur collaris) [Donati et al., 2007],
and orangutans (Pongo pygmeus) [Knott, 1998].
However, to our knowledge, only one report provided
the fatty acid composition of commonly consumed
foods by a wild primate species, the folivorous howler
monkeys (Alouatta palliata) [Chamberlain et al.,
1993]. Still, previous reports of the crude fat
consumed by primates have yielded interesting
results, particularly in comparison to human diets
[Milton, 1999]. Plant material commonly eaten by
orangutans varied from undetectable to 5% crude fat
on a dry matter (DM) basis; however, one seasonally
important food (Neesia sp. seed) contained a very high
concentration of lipids (46%) [Knott, 1998]. In a
community‐wide study of rainforest primates, plant
foods consumed by red‐tailed monkeys (Cercopithe-
cus ascanius), blue monkeys (Cercopithecus mitis),
gray‐cheeked mangabeys (Lophocebus albigena), and
chimpanzees (P. troglodytes) in Kibale National Park,
Uganda revealed the highest lipid intakes (6–8.5%) of
DM intake occurred during periods of fruit abun-
dance, which is as expected since fruits (which
include seeds) contain more lipid than leaves and
flowers. On average, lipids comprised 2.5–4% of DM
intake for these Cercopithecines [Conklin‐Brittain
et al., 1998]. Saki monkeys ate diets high in total lipid
(16.1%), chiefly because they are seed‐eaters, and
consume large quantities of lipid‐rich seeds daily
[Norconk and Conklin‐Brittain, 2004]. Mongoose
lemurs and collared lemurs had diets low in total
lipid (2%), probably due to their diet consisting
mainly of flowers, sugary fruits, and leaves
[Curtis, 2004; 4–5%; Donati et al., 2007].
Am. J. Primatol.
Modern human diets are higher in crude fat (15–
25% of DM) [Popovich et al., 1997; Wrangham
et al., 1998] than wild primate diets studied to date
aside from the seed‐eating saki monkeys [Conklin‐
Brittain et al., 1998]. In many populations worldwide,
more than 30% of energy intake is from crude fat
[WHO, 2003], and lipid intake for humans in
industrialized populations is estimated to compose
up to 40% of total energy intake [Jones et al., 1985].
These high levels of fat in the diet are thought to
contribute to diseases of public health concern
such as obesity, heart, and liver disease [USDA;
USHSS, 2010]. In the United States, it is recom-
mended that adults consume 20–35% of calories per
day as lipid [USDA; USHSS, 2010]. Estimates of
actual total lipid intake in American diets range from
34% [USDA; USHSS, 2010] to 42% of energy from
lipids [Lissner et al., 1987]. Humans acquire essen-
tial fatty acids from aquatic sources, meat, seeds,
legumes, fruits and vegetables, including some leafy
vegetables such as spinach, red leaf lettuce and
mustard [Simopoulos, 1991].
Mountain gorillas (Gorilla beringei) in Bwindi
Impenetrable National Park (BINP), Uganda con-
sume diets consisting almost entirely of plant tissues
[Rothman et al., 2006; Stanford & Nkurunungi, 2003;
Watts, 1984], and thus acquire fatty acids from
plants, though some fatty acids may be acquired from
intentionally or accidentally consumed rare foods
including ants or other insects [Ganas & Robbins,
2004]. Bwindi gorillas are opportunistic frugivores.
They mainly consume a diverse diet of leaves, pith,
stems, and fruit [Rothman et al., 2006] and depend
heavily on herbaceous vegetation during periods of
fruit scarcity [Stanford & Nkurunungi, 2003]. The
mountain gorillas studied previously in this same
location consumed a staple diet (97% of the annual
intake) that consisted mainly of herbaceous leaves,
as well as pith, the outer peel of herb stems
(later referred to as peel), fruit, and decaying wood
[Rothman et al., 2007]. The staple diet is rich in
crude protein (18%), fiber (43%), and non‐structural
carbohydrates (19%) [Rothman et al., 2007]. The
gorilla diet varies seasonally [Rothman et al., 2008].
These gorillas prioritize the consumption of non‐
protein energy (carbohydrates and fats) and consume
a high‐protein diet across seasons [Rothman
et al., 2011].
Here, we provide the first quantification of the
fatty acid composition of the wild diets of great apes,
which is based on our analyses of the fatty acid
composition and crude fat (EE) intake of feeds
comprising the staple diet of Bwindi mountain
gorillas. We also describe the fatty acid composition
of different plants and plant parts in the staple diet,
and discuss the possible effects this variation may
have on the total fatty acid intake. Finally, we
address ways that our results may inform mountain
gorilla feeding ecology and primate diets in general.
 Fatty Acids in Gorilla Diets / 3

METHODS sample was sealed in a labeled paper bag until

Foods that were part of the staple diet consumed
by mountain gorillas (Gorilla beringei) in BINP,
Uganda (0 530 –1 080 S and 29 350 –29 500 E) were
analyzed for their EE content and fatty acid
composition. This research adhered to the legal
requirements of the country in which the research
was conducted.
Staple Diet Determination
Mountain gorillas were studied over the course of
a year, quantifying their staple diet using behavioral
observations and measures of feed intake [Rothman
et al., 2007]. The staple diet was defined as any food
item that comprised 1% DM of intake annually. The
staple diet was comprised of 18 foods, with nine
different species of herbaceous leaves providing
60.5% of total dietary intake of DM, 14.3% from fruit,
6.6% from tree leaves, 6.3% from pith, 5.4% from peel,
and 4.0% from decaying wood [Rothman et al., 2007]
(Table I). The remaining 3% of the total diet was
composed of miscellaneous foods that were rarely
consumed, or consumed in negligible amounts
annually [Rothman et al., 2007].
Sample Collection
Plant items comprising 1% DM of the mountain
gorilla’s annual diet were collected fresh from BINP.
Once collected, samples were taken back to the field
station and dried at ambient temperature out of
direct sunlight at the field station (22°C) [Rothman
et al., 2007, 2008]. Following the drying process, each
ground to pass through a 1‐mm screen using a Wiley
Mill. Ground samples were then stored in a dry, dark
area until transported to the Nutritional Ecology
Laboratory at Hunter College, New York, USA and
stored in a dry, dark area until assayed [Rothman
et al., 2007, 2008]. In addition, because unsaturated
fatty acids are susceptible to oxidation [Czerkawski,
1967; Molloy et al., 1974; but see: Arvidsson et al.,
2009], on January 23, 2013, 1–3 g of each food species
was recollected in BINP and were immediately
placed in liquid nitrogen in 5 ml cryotubes for fatty
acid analysis.
Crude Fat Analysis
Percentage of EE was measured at Hunter
College, New York, USA using high‐temperature
petroleum ether extraction via an XT15 Fat Analyzer
[ANKOM, Macedon, New York], whereby the dried,
milled food samples of 0.45–0.55 g were boiled in
petroleum ether in filter bags at 90°C for 120 min
[AOCS, 2009]. After boiling, the samples were dried
in an oven at 90°C for 8 h and weighed. Assays were
calibrated with an 8.7% fat standard provided by
ANKOM. Three different samples of each of the
staple feeds, with the exception of four of the feeds,
due to limited sample, were analyzed. Only one
sample of Cyathea manniana pith, and Momordica
foetida herb leaves were available, and only two
were available for Maesa lanceolata fruit. While M.
foetida and Momordica pterocarpa are different
foods, they were combined by Rothman et al. [2007]
in behavioral observations because their phenotypic

TABLE I. Daily Intake and Crude Fat of Foods in the Staple Diet of Mountain Gorillas

Plant Plant part % Intakea % EEb

Basella alba Herbaceous leaves 7.8 1.3

Carduus Herbaceous leaves 4.2 3.4
Chrysophyllum albidum Fruit 3.0 19.0
Cyathea manniana Pith 2.2 1.2
Decaying wood Wood 4.0 0.0
Drypetes Fruit 0.9 1.1
Ipomea involucrata/Ipomea wightic Herbaceous leaves 8.8 2.1
Maesa lanceolata Fruit 1.8 28.4
Mimulopsis arborescens Pith 4.1 0.5
Mimulopsis solmsii Herbaceous leaves 7.1 1.4
Myrianthus holstii Fruit 8.6 3.3
Tree leaves 6.6 1.6
Momordica foetida/Momordica pterocarpac Herbaceous leaves 8.0 2.9
Triumfetta tomentosa Herbaceous leaves 5.4 2.1
Urera hypselodendron Herbaceous leaves 19.2 2.6
Peel 5.4 3.0
Total % intake of staple foods 97.1
a
The staple diet and % daily intake of each staple food is based on dry‐matter and was determined based on the wet weight intake by Rothman et al. [2007].
b
Ether Extract (measure of crude fat).
c
These herbaceous leaves were combined due to their similar phenotypes, making them difficult to differentiate [Rothman et al., 2007].

Am. J. Primatol.
4 / Reiner et al.
characteristics made it difficult to differentiate them
with a high level of accuracy. For this report, M.
foetida and M. pterocarpa were combined in all
calculated nutritional estimations of dietary intake,
EE, and fatty acid results. Similarly, Ipomea in-
volucrata and Ipomea wighti herb leaves were
combined in all calculated nutritional estimations
of dietary intake, EE, and fatty acid results due to
difficulty differentiating samples phenotypically dur-
ing behavioral observations. The fatty acids in the
Drypetes fruit had oxidized due to experiencing a
difference in handling method, so the fatty acid
results for that food were excluded from results and
analysis. However, this fruit was low in fat (1.1%) and
accounted for less than 1% of DM intake, and so its
exclusion is unlikely to affect the results.
Fatty Acid Analysis
The liquid nitrogen frozen samples were shipped
in a dry shipper (Arctic Express, Thermo Fisher
Scientific) to the Diagnostic Center for Population
and Animal Health at the University of Michigan,
USA where they were analyzed for their fatty acid
composition using gas–liquid chromatography on a
DM basis [IUPAC, 1997]. Samples were analyzed in
duplicate.
Crude Fat and Fatty Acid Intakes
The EE and fatty acid composition of the diet
were calculated using the weighted intake of each
staple food in its contribution to the total diet
[Rothman et al., 2007]. Using these intake data,
the total fatty acid composition of the diet and the
intake ratios for polyunsaturated fatty acids to
saturated fatty acids (SFA), n‐6:n‐3, and LA:ALA
were calculated.
When calculating the fatty acid intake from
feeds, defined as the sum of all FA obtained from the
lipid extract that are expressed as triacylglycerols, it
is necessary to correct for the weight of lipid extract
from non‐fat lipids (nonsaponifiable substances), so a
value of one was subtracted from the percentage of
ether extract (EE) to estimate the non‐fat compo-
nents that were extracted with ether [Palmquist &
Jenkins, 2003; Rothman et al., 2012]. To estimate
the metabolizable energy contributions of lipid to the
diet in relation to protein, fiber, and nonstructural
carbohydrates, the energetic contributions from
adjusted EE (9 kcal/g), crude protein (4 kcal/g), non‐
structural carbohydrates (4 kcal/g), and neutral
detergent fiber (3 kcal/g multiplied by a period and
age–sex specific digestibility coefficient [Rothman
et al., 2008]) were added together [Conklin‐Brittain
et al., 2006; Rothman et al., 2012]. The percentages of
dietary crude protein, non‐structural carbohydrates,
and neutral detergent fiber were previously calculat-
ed by Rothman et al. [2007].
Am. J. Primatol.
This research adhered to the American Society of
Primatologists principles for the ethical treatment of
primates.
RESULTS
Of the staple foods, only decaying wood contained
undetectable levels of EE. Crude fat comprised 3.1%
of the total dietary intake for these gorillas (DM),
about 14% of the total energy intake, and about 22%
of the non‐protein energy (Table I). Fruit contained
13.0% (SD 13%) fat on average, and provided about
44.9% of all FA in the staple diet with the other 55.1%
coming from other staple vegetative parts (herba-
ceous leaves, peel, pith, tree leaves). Herbaceous
leaves were low in fat (2.3%, SD 0.8%) but supplied
44.8% of dietary FA due to their high proportional
intake (60.5% of estimated intake). Peel (5.3%), tree
leaves (3.5%), and pith (1.5%) supplied small
amounts of FA in the staple diet (Fig. 1).
SFAs, monounsaturated fatty acids (MUFA), and
PUFA are the three main classes of fatty acids. Total
fatty acids were comprised of 32.4% SFA, 12.5%
MUFA, and 54.6% PUFA (with about 0.21% uniden-
tified fatty acids) (Fig. 1; Table II). Of the total fatty
acid composition of the diet, 47.4% of SFA were
supplied by fruit and 41.1% from herbaceous leaves.
Peel, tree leaves, and pith were all very low
contributors to total diet SFA (each <6% of total
FA) compared to herbaceous leaves and fruit. Fruit
provided most of the MUFA in the diet (54.5%)
followed by herbaceous leaves (36.4%). Herbaceous
leaves provided the most PUFA in the diet (49.2%)
followed by fruit (41.1%). The total of all n‐6 PUFA in
the diet was 32.2%
SD 10.0%, while the n‐3 PUFA
composed 22.4%
SD 8.6%. The most prevalent SFA
(>4% of total FA) were palmitic acid C16:0 (23.9%)
and stearic acid C18:0 (5.7%) (Table II). The most
prevalent MUFA and PUFA in the diet were oleic
100 n-6 Polyunsaturated Fay Acids
n-3 Polyunsaturated Fay Acids
90
Monunsaturated Fay Acids
80 Saturated Fay Acids
70
60
50
40
30
20
10
0
Total Diet Herbaceous Fruit Peel Tree Leaves Pith
Leaves
Fig. 1. Percentage of fatty acids supplied in mountain gorilla
(Gorilla beringei) staple diet by plant part.
 Fatty Acids in Gorilla Diets / 5

TABLE II. Fatty Acid Composition of Wild Mountain DPA (C22:5n‐3), and DHA (C22:6n‐3) (DHA); howev-
Gorilla Diets (G. beringei) er, the total percentage of these long‐chain PUFA was
low (1%).
g FA/100g
Fatty acid Common name gorilla dietd
DISCUSSION
Saturated fatty acids
C6:0 Capronic 0.000 Although the macronutrients of wild primate
C8:0 Caprylic 0.000 diets have been studied, very little is known about
C10:0 Capric 0.000 micronutrients such as fatty acids, amino acids, and
C12:0 Lauric 0.012 minerals [but see: Chamberlain et al., 1993; Milton &
C14:0 Myristic 0.051 Dintzis, 1981; Nagy & Milton, 1979a, b]. Our study
C16:0 Palmitic 0.732 examines the sources of dietary fatty acids for
C18:0 Stearic 0.175 mountain gorillas and estimates the amounts of total
C20:0 Arachidic 0.001
fat and dietary fatty acid intake by wild mountain
C22:0 Behenic 0.000
C24:0 Lignoceric 0.000 gorillas. A limitation of the study is that only the
Total SFAa 0.992 staple portion of the gorilla diet was analyzed;
Monounsaturated fatty acids however, those staple foods were estimated to
C12:1n1c n/a 0.011 account for 97% of food intake.
C14:1n5c Myristoleic 0.012 The wild mountain gorilla diet is low in total lipid
C16:1n‐7c Palmitoleic 0.047 (3.1% of DM) in comparison with the diet of most
C16:1n7t Palmitelaidic 0.014 modern humans [Institute of Medicine (US) &
C18:1n‐7c Vaccenic 0.013 Institute of Medicine (US), 2005; Lissner et al.,
C18:1n‐9c Oleic 0.259 1987; USHSS, 2010; WHO, 2003], and similar to the
C18:1n‐9t Elaidic 0.004
fat contents in the diets of primates that are
C20:1n‐9c n/a 0.000
C22:1n‐9c Erucic 0.004 folivores–frugivores [Conklin‐Brittain et al., 1998;
C24:1n‐9c Nervonic 0.000 Curtis, 2004; Donati et al., 2007] though lower in fat
Total MUFAb 0.381 than diets of seed eating primates [Norconk and
Polyunsaturated n‐3 fatty acids Conklin‐Brittain, 2004]. Crude fat composes just 14%
C18:3n‐3c Alpha‐linolenic 0.651 of the total energy intake in the diets of mountain
C20:3n3c n/a 0.018 gorillas, and accounts for 22% of the non‐protein
C20:5n‐3c Eicosapentaenoic 0.004 energy ingested. Fruit and herbaceous leaves provid-
C22:5n‐3c Docosapentaenoic 0.003 ed the majority of essential fatty acids in the diet of
C22:6n‐3c Docosahexaenoic 0.010 mountain gorillas, accounting for 89.7% of the fatty
Total n‐3 PUFAc 0.686
acid intake while comprising 74.8% of total DM
Polyunsaturated n‐6 fatty acids
C18:2n‐6c Linoleic 0.927 intake. Although fruit accounted for only 14.3% of
C18:3n‐6c Gamma linolenic 0.034 dietary intake, fruit provided 44.9% of the fatty
C20:4n‐6c Arachidonic 0.001 acid intake and a majority of the intake of the n‐6
C22:4n‐6c Adrenic 0.024 essential fatty acids LA. Thus, in addition to
Total n‐6 PUFA 0.987 providing other important nutrients, fruit contribut-
Total PUFA 1.673 ed to total dietary fat intake and intake of essential
a
Saturated fatty acid (does not contain any double carbon bonds).
fatty acids out of proportion to its overall intake.
b
Monounsaturated fatty acid (one double carbon bond). Herbaceous leaves were low in fat, but because they
c
d
Polyunsaturated fatty acid (2 double carbon bonds)]. comprise 60.5% of the total dietary intake they
Of the staple diet; based on dry‐matter, and determined by Rothman et al.
[2007]. provide 44.8% of the total fatty acids and a majority of
the n‐3 essential fatty acids. Only one of the staple
feed items is a tree leaf (Myrianthus holstii), and it
contributed 6.6% of the staple diet and provided 3.5%
acid C18:1n9c (8.5%), LA (30.3%), and ALA (21.2%) of the total fatty acids. Other plant parts contributed
(Table II). LA and ALA were the most prevalent small portions of fatty acids to the total composition of
PUFA in the diet and were detected in all of the EE the diet (peel 5.3%, tree leaf 3.5%, pith 1.5%). Thus,
containing staple feeds though their ratios differed by fruits (due to their higher lipid content) and herba-
diet item. Fruit provided the majority of the n‐6 ceous leaves (due to the quantity consumed) are the
essential fatty acid (LA) in the diet, making up 52.5% main contributors of lipid and essential fatty acids in
of the total LA. The majority of the n‐3 essential fatty the diet.
acid (ALA) in the diet was provided by herbaceous Together, LA and ALA comprised 51.5% of the
leaves (67.9%). The ratio of LA:ALA in the total diet total FA consumed by the gorillas. Consistent with
was 1.44. the finding that herbaceous leaves provided more
Fruit provided the most AA (C20:4n‐6), while ALA while fruits provided higher amounts of LA,
herbaceous leaves provided the most EPA (C20:5n‐3), herbaceous leaves provided the most EPA, DPA, and

Am. J. Primatol.
6 / Reiner et al.
DHA, which are long‐chain derivatives of ALA, while
fruit provided the most AA, a long‐chain derivative of
LA. Dietary deficiencies in essential fatty acids and
their long‐chain derivatives are linked to neurologi-
cal abnormalities in humans and other mammals
[Bjerve et al., 1987; Holman, 1998; Moriguchi &
Salem, 2003]. While AA, EPA, DPA, and DHA are not
essential fatty acids in the strict sense because
primates can derive them endogenously from shorter
chain fatty acids (LA and ALA), these long‐chain
PUFA are essential for proper brain, retinal, neural,
and immune function [Moriguchi & Salem, 2003;
Uauy & Dangour, 2006]. The conversion of shorter
chain fatty acids to AA and EPA readily occurs
[Brenna et al., 2009; Su et al., 1999], while the
conversion to DHA is only 1% in humans [Brenna
et al., 2009] and likely similar in most primates.
Despite the finding that the total percentage of AA,
EPA, DPA, and DHA from all staple diet foods was
only 0.6%, the high levels of the essential fatty acids
LA and ALA in the diet suggest that the longer chain
PUFAs are not limiting factors and wild mountain
gorilla diet is not deficient in long‐chain PUFAs.
Different plant parts will often differ in nutrient
composition [Rothman et al., 2006]. As such, we
expected that different parts of a single plant species
would vary in fatty acids composition. Two plant
species in the mountain gorilla staple diet were each
represented by two different plant parts. Myrian-
thus holstii fruit and tree leaves are consumed
regularly, in addition to Urera hypselodendron herb
leaves and peel. Consistent with the general differ-
ences between fruits and leaves, M. holstii fruit
contained more EE than the tree leaves (3.3% vs.
1.6%) and had a higher LA:ALA ratio (1.0 vs. 0.36)
and a lower PUFA:SFA ratio (1.2 vs. 2.4). Despite
fairly similar levels of intake (Table I) M. holstii fruit
contributed a larger percentage of fatty acid than M.
holstii tree leaves to the total diet (fruit 9.3% and
tree leaves 3.5%). Urera hypselodendron herb leaves
contained 2.6% EE which was similar to the value for
the peel (3.0% EE). However, U. hypselodendron
herb leaves and peel differed in LA:ALA ratio (0.73
vs. 3.49) and PUFA:SFA ratio (2.35 vs. 1.67). The
peel from U. hypselodendron contributed 5.3% of the
fatty acid intake, in proportion with its value for DM
intake, while the leaves from this plant species
contributed slightly lower to fatty acid intake
relative to DM intake (16.3% of fatty acids vs.
19.2%) due to the lower lipid content. The leaves
were still a more important contributor to fatty acid
intake.
Milligan et al. [2008] investigated the fatty
acid composition of milks from five wild anthropoids
(A. palliata, Callithrix jacchus, G. beringei, Leonto-
pithecus rosalia, Macaca sinica) and found that the
folivorous species (A. palliata and G. beringei)
produced milk with higher proportions of LA than
other three anthropoids studied. We found LA was
Am. J. Primatol.
one of the most predominant fatty acids in Bwindi
mountain gorilla diets, representing 30.3% of the
total fatty acids in the diet. It is possible that LA is in
higher concentration in mountain gorilla diets
compared to the wild diets of other primate species,
though more studies of dietary fatty acids content are
needed to test this hypothesis. Milligan et al. [2008]
noted that mountain gorillas were unique among
anthropoids in the high proportion of AA in their
milk. We found very low amounts (0.023% total FA) of
AA in the diet of Bwindi mountain gorillas. Because
dietary LA can be converted to AA after it is
consumed, the relatively high AA levels observed in
mountain gorilla milk could result from a high
dietary LA intake. The milk samples in this study
were collected from mountain gorillas in the Virunga
region, where gorillas eat different diets [Rothman
et al., 2007]; however, the diet results of this study are
consistent with the hypothesis that the high levels of
AA in mountain gorilla milk is related to dietary fatty
acid composition.
Of total fatty acids in the mountain gorilla staple
diet, about an eighth were MUFA (12.5%), a half were
PUFA (54.6%), and a third were SFA (32.4%). The
intake ranges recommended for all human popula-
tions for SFA is 10% of total energy and 6–11% for
total PUFA [WHO, 2008]. The ratio of PUFA:SFA
consumed by mountain gorillas was 1.69. The PUFA:
SFA ratio of some major diet components of wild
howler monkeys was 0.85 [Chamberlain et al., 1993],
and the recommendation for humans varies from 0.6
to 1.1 (6%–11% PUFA, <10% SFA) [WHO, 2003].
Thus the PUFA to SFA ratio for the mountain gorilla
diet falls above the high end of the range recom-
mended for humans. The diets of many human
populations are composed of SFA in amounts that
exceed nutritional recommendations for optimal
health. While gorillas consume a high proportion of
their dietary fat as SFA, the total energy contributed
by SFA is only about 3.2%, so it is well below the
maximum recommendation for humans. Reconstruc-
tions of pre‐agricultural paleolithic diets estimate
that humans and livestock such as cattle consume
diets with an equilibrant ratio (1:1) of total n‐6:n‐3
PUFA [Wathes et al., 2007]. Currently, humans
consume these FA in ratios that range from 10:1 to
25:1 in populations dependent on globalized and
industrial agriculture [Wathes et al., 2007], which
may have severe health consequences related to
obesity and heart disease [USDA; USHSS, 2010]. The
n‐6:n‐3 PUFA ratio of the gorilla staple diet was
found to be 1.44. Thus wild mountain gorillas are
consuming a ratio of n‐6:n‐3 PUFA closer to
agricultural paleolithic diets. This very low fat diet
in wild gorillas also has implications for feeding
gorillas in captivity and zoo managers should
consider the extent to which emulating wild diets
might prevent diet related illnesses [Crissey &
Pribyl, 1997; Grant et al., 2002].

ACKNOWLEDGMENTS
We thank the field assistants of the Institute of
Tropical Forest Conservation for their hard work in
the field. We also thank Hillary Musinguzi, Pontius
Ezuma and Simplicious Gessa for logistical support
in the field. We thank Jenny Paltan and Scott
Williams for laboratory assistance. Peter Van Soest
and Herman Pontzer provided important suggestions
on a draft of this manuscript. We are grateful to the
two reviewers who provided feedback that substan-
tially improved the manuscript with respect to our
fatty acid drying methods from the submitted draft.
The Uganda Wildlife Authority and the Uganda
National Council for Science and Technology gave us
permission for this research.
REFERENCES
Anderson BM, Ma DWL. 2009. Are all n‐3 polyunsaturated
fatty acids created equal? Lipids Health Dis 8:33.
AOCS. 2009. Official Procedure Am 5‐04. Rapid determination
of oil/fat utilizing high temperature solvent extraction.
Available online at: http://www.ankom.com/media/documents/
CrudeFat_0504_013009.pdf.
Arvidsson K, Gustavsson AM, Martinsson K. 2009. Fatty acids
in forages: a comparison of different pre‐treatments prior to
analysis. An Feed Sci Tech 151:143–152.
Bjerve K, Mostad I, Thoresen L. 1987. Alpha‐linolenic acid
deficiency in patients on long‐term gastric tube feeding;
estimation of linolenic acid and long chain unsaturated n‐3
fatty acid requirements in man. Am J Clin Nutr 45:66–77.
Brenna JT, Salem N Jr, Sinclair AJ, Cunnane SC. 2009.
a‐Linolenic acid supplementation and conversion to n‐3
long‐chain polyunsaturated fatty acids in humans. Prosta-
glandins Leukot Essent Fatty Acids 80:85–91.
Chamberlain J, Nelson G, Milton K. 1993. Fatty acid profiles of
major food sources of howler monkeys (Alouatta palliata) in
the neotropics. Cell Mol Life Sci 49:820–824.
Chapkin RS, Kim W, Lupton JR, McMurray DN. 2009. Dietary
docosahexaenoic acid and eicosapentaenoic acid: emerging
mediators of inflammation. Prostaglandins Leukot Essent
Fatty Acids 81:187–191.
Conklin‐Brittain N, Wrangham R, Hunt K. 1998. Dietary
response of chimpanzees and cercopithecines to seasonal
variation in fruit abundance. II. Macronutrients. Int J
Primatol 19:971–998.
Conklin‐Brittain N, Knott C, Wrangham R. 2006. Energy
intake by wild chimpanzees and orangutans: methodological
considerations and a preliminary comparison. In: Hohmann
G, Robbins MM, Boesch C, editors. Feeding ecology in apes
and other primates: ecological, physical, and behavioral
aspects. Cambridge: Cambridge University Press. p 445–
472.
Conner WE. 2000. Importance of n‐3 fatty acids in health and
disease. Am J Clin Nutr 71:171S–175S.
Crawford MA. 1992. The role of dietary fatty acids in biology:
their place in the evolution of the human brain. Nutr Rev
50:3–11.
Crawford MA, Bloom M, Broadhurst CL, Schmidt WF,
Cunnane SC. 1999. Evidence for unique function of
docosahexaenoic acid during the evolution of the modern
human brain. Lipids 34:S39–S47.
Crissey SD, Pribyl LS. 1997. Utilizing wild foraging ecology
information to provide captive primates with an appropriate
diet. Proc Nutr Soc 56:1083–1094.
Curtis DJ. 2004. Diet and nutrition in wild mongoose lemurs
(Eulemur mongoz) and their implications for the evolution of
Fatty Acids in Gorilla Diets / 7
female dominance and small group size in lemurs. Am J Phys
Anthropol 124:234–247.
Czerkawski JW. 1967. Effect of storage on the fatty acids of
dried ryegrass. Br J Nutr 21:599–608.
Donati G, Bollen A, Borgognini‐Tarli SM, Ganzhorn JU. 2007.
Feeding over the 24‐h cycle: dietary flexibility of cathemeral
collared lemurs (Eulemur collaris). Behav Ecol Sociobiol
61:1237–1251.
Ganas J, Robbins MM. 2004. Intrapopulation differences in ant
eating in the mountain gorillas of Bwindi Impenetrable
National Park, Uganda. Primates 45:275–278.
Grant JB, Brown DL, Dierenfeld ES. 2002. Essential fatty acid
profiles differ across diets and browse of black rhinoceros. J
Wildl Dis 38:132–142.
Holman RT. 1998. The slow discovery of the importance of
omega 3 essential fatty acids in human health. J Nutr
128:427S–433S.
Institute of Medicine (U.S.). 2005. Dietary reference intakes for
energy, carbohydrate, fiber, fat, fatty acids, cholesterol,
protein, and amino acids. Washington, DC: National
Academies Press.
IUPAC. 1997. Compendium of chemical terminology: IUPAC
recommendations. Compiled by McNaught AD, Wilkinson A,
editors. Oxford: Blackwell Science.
Jones P, Pencharz P, Clandinin M. 1985. Whole body oxidation
of dietary fatty acids: implications for energy utilization. Am
J Clin Nutr 42:769–777.
Knott C. 1998. Changes in orangutan caloric intake, energy
balance, and ketones in response to fluctuating fruit
availability. Int J Primatol 19:1061–1079.
Lissner L, Levitsky DA, Strupp BJ, Kalkwarf HJ, Roe DA.
1987. Dietary fat and the regulation of energy intake in
human subjects. Am J Clin Nutr 46:886–892.
Milligan L, Rappoport S, Cranfield M, et al. 2008. Fatty acid
composition of wild anthropoid primate milks. Comp
Biochem Physiol Biochem B Mol Biol 149:74–82.
Milton K. 1999. Nutritional characteristics of wild primate
foods: do the diets of our closest living relatives have lessons
for us? Nutrition 15:488–498.
Milton K, Dintzis F. 1981. Nitrogen‐to‐protein conversion
factors for tropical tree samples. Biotropica 13:177–181.
Molloy LF, Giltrap DJ, Collie TW, Metson AJ. 1974. Degrada-
tion of higher fatty acids in perennial ryegrass and white
clover following drying and storage. J Sci Food and Agri
25:595–606.
Moriguchi T, Salem N Jr. 2003. Recovery of brain docosahex-
aenoate leads to recovery of spatial task performance.
J Neurochem 87:297–309.
Nagy KA, Milton K. 1979a. Aspects of dietary quality, nutrient
assimilation and water balance in wild howler monkeys
(Alouatta palliata). Oecologia 39:249–258.
Nagy KA, Milton K. 1979b. Energy metabolism and food
consumption by wild howler monkeys (Alouatta Palliata).
Ecology 60:475–480.
Norconk MA, Conklin‐Brittain NL. 2004. Variation on frugi-
vory: the diet of Venezuelan white‐faced sakis. Int J Primatol
25:1–26.
Palmquist D, Jenkins T. 2003. Challenges with fats and fatty
acid methods. J Anim Sci 81:3250–3254.
Pereira SL, Leonard AE, Mukerji P. 2003. Recent advances in
the study of fatty acid desaturases from animals and lower
eukaryotes. Prostaglandins Leukot Essent Fatty Acids
68:97–106.
Popovich DG, Jenkins DJA, Kendall CWC, Dierenfeld ES,
Carroll RW, Tariq N, Vidgen E. 1997. The western lowland
gorilla diet has implications for the health of humans and
other hominoids. J Nutr 127:2000–2005.
Rothman JM, Dierenfeld ES, Molina DO, et al. 2006.
Nutritional chemistry of foods eaten by gorillas in Bwindi
Impenetrable National Park, Uganda. Am J Primatol 68:
675–691.
Am. J. Primatol.
8 / Reiner et al.
Rothman JM, Plumptre AJ, Dierenfeld ES, Pell AN. 2007.
Nutritional composition of the diet of the gorilla (Gorilla
beringei): a comparison between two montane habitats.
J Trop Ecol 23:673–682.
Rothman JM, Dierenfeld ES, Hintz HF, Pell AN. 2008.
Nutritional quality of gorilla diets: consequences of age,
sex, and season. Oecologia 155:111–122.
Rothman JM, Raubenheimer D, Chapman CA. 2011. Nutri-
tional geometry: gorillas prioritize non‐protein energy while
consuming surplus protein. Biol Lett 7:847–849.
Rothman JM, Chapman CA, Van Soest PJ. 2012. Methods in
primate nutritional ecology: a user’s guide. Int J Primatol
33:542–566.
Simopoulos AP. 1991. Omega‐3 fatty acids in health and
disease and in growth and development. Am J Clin Nutr
54:438–463.
Stanford CB, Nkurunungi JB. 2003. Behavioral ecology of
sympatric chimpanzees and gorillas in Bwindi Impenetrable
National Park, Uganda: diet. Int J Primatol 24:901–
918.
Su H‐M, Corso TN, Nathanielsz PW, Brenna TJ. 1999. Linoleic
acid kinetics and conversion to arachidonic acid in the
pregnant and fetal baboon. J Lipid Res 40:1304–1311.
Uauy R, Dangour AD. 2006. Nutrition in brain development
and aging: role of essential fatty acids. Nutr Rev 64:S24–
S33.
Am. J. Primatol.
U.S. Department of Agriculture and U.S. Department of Health
and Human Services. 2010. Dietary guidelines for Ameri-
cans, 2010. 7th edition. Washington, DC: U.S. Government
Printing Office.
Wainwright P, Ward G. 1997. Early nutrition and behavior:
a conceptual framework for critical analysis of research.
In: Dobbing J, editor. Developing brain and behaviour: the
role of lipids in infant formula. San Diego: Academic Press.
p 387–414.
Wathes C, Abayasekara R, Aitken R. 2007. Polyunsaturated
fatty acids in male and female reproduction. Biol Reprod
77:190–201.
Watts DP. 1984. Composition and variability of mountain
gorilla diets in the Central Virungas. Am J Primatol 7:323–
356.
World Health Organization. 2003. Joint WHO/FAO expert
consultation on diet, nutrition and the prevention of chronic
diseases. WHO Technical Report Series 916. Geneva,
Switzerland: WHO.
World Health Organization. 2008. WHO|Joint FAO/WHO
expert consultation on fats and fatty acids in human
nutrition. Geneva, Switzerland: WHO.
Wrangham R, Conklin‐Brittain N, Hunt K. 1998. Dietary
response of chimpanzees and cercopithecines to seasonal
variation in fruit abundance. I. Antifeedants. Int J Primatol
19:949–970.