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Comparative use of color vision for frugivory by sympatric species of platyrrhines.

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American Journal of Primatology 67:399–409 (2005)
Research Article
Comparative Use of Color Vision for Frugivory by
Sympatric Species of Platyrrhines
Centro de Investigaciones en Ecosistemas, Universidad Nacional Autónoma de México,
Morelia, Michoacán, Me´xico
Escuela de Biologı´a, Universidad de Costa Rica, San José, Costa Rica
Department of Anatomy, University of Hong Kong, Hong Kong, PR China
Ateles spp. and Alouatta spp. are often sympatric, and although they are
mainly frugivorous and folivorous, respectively, they consume some of
the same fruit species. However, they differ in terms of color vision,
which is thought to be important for fruit detection. Alouatta spp. have
routine trichromatic color vision, while Ateles spp. presents the classic
polymorphism of platyrrhines: heterozygous females have trichromatic
color vision, and males and homozygous females have dichromatic vision.
Given these perceptual differences, one might expect Alouatta spp. to
consume more reddish fruits than Ateles spp., since trichromats have an
advantage for detecting fruits of that hue. Furthermore, since Ateles spp.
have up to six different color vision phenotypes, as do most other
platyrrhines, they might be expected to include fruits with a wider variety
of hues in their diet than Alouatta spp. To test these hypotheses we
studied the fruit foraging behavior of sympatric Alouatta palliata and
Ateles geoffroyi in Costa Rica, and modeled the detectability of fruit via
the various color vision phenotypes in these primates. We found little
similarity in fruit diet between these two species (Morisita 5 0.086).
Furthermore, despite its polymorphism, A. geoffroyi consumed more
reddish fruits than A. palliata, which consumed more greenish fruits.
Our modeling results suggest that most fruit species included in the diet
of A. geoffroyi can be discriminated by most color vision phenotypes
present in the population. These findings show that the effect of
polymorphism in platyrrhines on fruit detection may not be a
disadvantage for frugivory. We suggest that routine trichromacy may
be advantageous for other foraging tasks, such as feeding on young
leaves. Am. J. Primatol. 67:399–409, 2005. c 2005 Wiley-Liss, Inc.
Contract grant sponsor: Research Grants Council of Hong Kong; Contract grant number: 7241/
97M; Contract grant sponsor: National Geographic Society; Contract grant number: 6584-99;
Contract grant sponsor: Marenco Beach and Rainforest Lodge.
Correspondence to: Kathryn E. Stoner, Centro de Investigaciones en Ecosistemas, Universidad
Nacional Autónoma de México, Apartado Postal 27-3 (Xangari), Morelia, Michoacán, México 58980.
Received 30 November 2004; revised 16 May 2005; revision accepted 20 May 2005
DOI 10.1002/ajp.20195
Published online in Wiley InterScience (
r 2005 Wiley-Liss, Inc.
400 / Stoner et al.
Key words: visual ecology; Ateles geoffroyi; Alouatta palliata; color
vision; frugivory; Costa Rica
Spider (Ateles spp.) and howler (Alouatta spp.) monkeys occur sympatrically
in most of their range [Emmons, 1990; Reid, 1997]. The former genus has been
considered a fruit specialist, with fruits accounting for 55–90% of their overall
diet [van Roosmalen, 1985; Chapman, 1987; Symington, 1987; Riba-Hernández &
Stoner, 2005]. In contrast, howler monkeys have been regarded as folivorousfrugivorous monkeys [Milton, 1980; Guillotin et al., 1994], with fruits contributing to 25–50% of their overall diet [Milton, 1980; Estrada, 1984; Estrada &
Coates-Estrada, 1984; Guillotin et al., 1994; Stoner, 1996]. While Alouatta spp.
consumes the same fruit species as Ateles spp. during certain periods of the year
[Chapman, 1988; Guillotin et al., 1994; Stevenson et al., 2000; Regan et al., 2001],
paradoxically, the genera have different color-vision capacities. Ateles spp. shares
the common M/L opsin polymorphism with most other platyrrhines, and has
dichromatic and trichromatic individuals in the same population. Homozygous
individuals (all males and about one-third of females) are dichromatic, while
heterozygous individuals (about two-thirds of females) are trichromatic.
Furthermore, platyrrhines with three alleles at the M/L locus can have up to
six different phenotypes in a population [Mollon et al., 1984]. In contrast,
Alouatta spp. is (thus far) the only platyrrhine genus to show routine
trichromacy. All individuals in this genus (both male and females) have three
opsins tuned at approximately 430 nm, 530 nm, and 560 nm [Jacobs et al., 1996].
It has been hypothesized that trichromacy is an adaptation made by primates
to detect food items [Allen, 1879; Polyak, 1957], with the selective advantage lying
in the ability to detect either red/orange fruits or young reddish leaves against
mature green foliage [Dominy & Lucas, 2001; Osorio & Vorobyev, 1996; Lucas
et al., 2005]. Foods with this pigmentation will show no color contrast for
individuals possessing dichromatic color vision, and hence will not be detectable
[Regan et al., 2001]. However, dichromatic vision could be advantageous for
detecting hue-camouflaged fruits, since this kind of vision reduces ‘‘chromatic
noise’’ and improves the detection of contours [Regan et al., 2001], which results
in breaking camouflage patterns [Morgan et al., 1992]. Accordingly, primates with
trichromatic vision would be expected to include more red/orange fruits in their
diets, while polymorphic species would show less reliance on foods with such color
hues (i.e., green fruits would be more important in their diet).
Several studies have evaluated the diet of Ateles spp. and Alouatta spp. and
shown that they consume some of the same fruit species; however, in most of
these investigations no quantitative comparisons of fruit color were made
[Chapman, 1987, 1988; Guillotin et al., 1994; Stevenson et al., 2000; Stoner, 1996;
Symington, 1988]. In a study in French Guiana, Guillotin et al. [1994] found no
differences between Alouatta seniculus and Ateles paniscus in the color of fruits
eaten, based on samples of stomach contents. Most fruits in the diet of both
species were orange-yellow or varying intensities of red, and fruits with green and
brown coloration were the least consumed. In another study in French Guiana,
Julliot [1996] found that A. seniculus preferred yellow and green fruits to red
fruits. Finally, a study that quantitatively compared color of fruits consumed by
Alouatta seniculus and Ateles paniscus in French Guiana [Regan et al., 2001]
found that trichromatic A. seniculus included less reddish fruits in their diet
Am. J. Primatol. DOI 10.1002/ajp
Use of Color Vision for Frugivory / 401
compared to polymorphic A. paniscus (2.9% vs. 14.6%). Nevertheless, it is unclear
how many fruit species were sampled for each monkey species, the fruits were not
necessarily collected while the animals were foraging, and the relative importance
of the fruit species in the diet is unknown. In summary, the current evidence is
insufficient to draw any conclusions relevant to evolutionary patterns in color
vision in platyrrhine monkeys. Systematic parallel studies of fruit foraging in
sympatric Alouatta spp. and Ateles spp. that document fruit species overlap in diet
and the relative importance of fruit species must be conducted before we can
quantify the importance of color differences in fruits that are consumed.
Accordingly, that was the aim of the present study. We studied Ateles geoffroyi
and Alouatta palliata over an annual period to 1) determine whether sympatric
A. palliata consumes more reddish fruits than A. geoffroyi, since trichromats have
an advantage for detecting fruits of this hue; and 2) to identify whether
A. geoffroyi includes a wider variety of fruit hues in its diet than A. palliata. Since
previous studies (cited above) have shown that Ateles spp. and Alouatta spp.
prefer ripe fruit, and many of the eaten species turn red or orange when ripe
[Sumner & Mollon, 2000b], we expected the fully trichromatic A. palliata to
restrict its fruit foraging to hues in this range because they have the ability to
distinguish and select these fruits. In contrast, A. geoffroyi, which includes some
dichromatic individuals that cannot distinguish red hues, would be anticipated to
randomly consume a greater variety of colors because they do not all have the
capacity to distinguish the critical hues. We employed modeling techniques to see
if differences in fruit choice were logical with respect to the capacity to
discriminate color afforded by the different possible opsin combinations.
Study Site and Species
The study was conducted at the Punta Rio Claro Wildlife Refuge (81390 N,
831440 E) located in the Osa Peninsula, southwestern Costa Rica. This area is
classified as tropical humid forest [Holdridge et al., 1971]. The mean annual
rainfall is approximately 3,000 mm, with a marked dry season from December–
April [Hartshorn, 1983]. Ateles geoffroyi Kuhl 1820 [Rylands et al., 2000], one of
four species in the genus, is distributed from Tamaulipas and Jalisco, Mexico, on
both coasts, to Oaxaca and southeastern Panama [Reid, 1997]. Alouatta palliata
Gray 1849 [Rylands et al., 2000] is distributed from Veracruz, Oaxaca, Chiapas,
and Tabasco, Mexico, to Colombia and west Ecuador [Reid, 1997].
Although a previous study reported the presence of only two alleles in Ateles
[Jacobs & Deegan, 2001], three alleles were found in our study population.
Sequencing of opsin genes was done by isolating DNA from fecal samples obtained
from 10 females and two males. This indicated the presence of 535 nm, 550 nm,
and 562 nm M/L alleles (W.-H. Li, personal communication) in the population.
Since these three alleles were found in our study population, even though we do
not know their exact frequency, we assume they are represented by a normal
distribution of color vision phenotypes among individuals. Therefore, we
performed our analyses using six possible phenotypes. In agreement with Jacobs
and Deegan [2001], Hiramatsu et al. (this volume) found only two alleles for
Ateles, but for Cebus they found three alleles in one population and two in
another, which indicates that in some platyrrhine monkeys the number of alleles
may vary between populations. The sequence of opsin genes in Alouatta spp.
shows that this species is routinely trichromatic [Jacobs et al., 1996].
Am. J. Primatol. DOI 10.1002/ajp
402 / Stoner et al.
Foraging Data and Analyses
The foraging behavior of one troop of A. geoffroyi containing five adult males,
10 adult females, six infants, and nine juveniles, and two troops of A. palliata (one
containing eight adult females, three adult males, four infants, and three
juveniles, and another containing four adult males, nine adult females, two
infants, and five juveniles) were studied from May 1999 through May 2000. We
used 2-min focal animal observations to collect data on fruit consumption by both
monkey species [Altmann, 1974]. To facilitate the location of A. palliata troops,
one female in each of the two troops was marked with a radiocollar transmitter
(AVM; Instruments Company, Colfax, CA). A permit was obtained from the Costa
Rican Wildlife Department to immobilize the monkeys, and we tranquilized the
animals with a licensed veterinarian present at all times. The tranquilized
animals were captured in hammock-type mesh nets, and no injuries occurred.
Data were collected 2 days per week for each species, from 0600 to 1800 hr. All
individuals were identified as to sex and age class, and the focal animals were
randomly changed after each 2-min observation.
Fruits were considered consumed when the monkeys bit into the fruit more
than twice and swallowed either pulp and seed(s) or the entire fruit. Samples of
fruits were collected when the monkeys mishandled fruit pieces before they fully
consumed it. When fruit samples were not obtained during monkey foraging
bouts, we returned to the same tree the following day and used a telescopic tree
pruner to collect samples from the branch where foraging was observed. The
percentage of fruits, leaves, and flowers in the overall diet was calculated as the
total time spent consuming items in each category divided by the overall foraging
time. The fruit species consumption time was calculated as the time dedicated to
consuming a fruit species divided by the total time spent consuming all fruit
species. The diet overlap of fruit was then calculated using the Morisita index
[Horn, 1966], given by:
2xi yi
x2i þ y2i
where xi is the proportion of the fruit species i in the diet of A. palliata (x), and yi
is the proportion of the same fruit species in the diet of A. geoffroyi (y). The sum
included all fruit species consumed by both primate species. The values of this
index range from 0 (no dietary overlap) to 1 (dietary overlap). Diet overlap was
calculated for the annual fruit diet.
Fruit and Leaf Background: Measurement of Reflectance Spectra
We followed Riba-Hernández et al. (this volume) in recording the fruit
and leaf background reflectance of fruit species consumed by both monkeys.
Reflectance spectra were recorded from the fruit coverings of fruits
consumed by the monkeys. To describe the background colors against which
fruits were seen, we collected two mature leaves surrounding the fruits where
monkeys were feeding. Only the upper surfaces of these leaves (which we
presumed the monkeys were observing) were recorded. Reflectance spectra
were recorded in the field using a portable field kit [Lucas et al., 2001]. Samples
were placed in a purpose-built chamber connected to a spectrometer
with illumination provided by a 12 V 3100k tungsten halogen lamp (LS-1; Ocean
Optics, Palo Alto, CA). The spectrometer was connected to a laptop portable
Am. J. Primatol. DOI 10.1002/ajp
Use of Color Vision for Frugivory / 403
computer via a PCMCIA card (DAQCard1200; National Instruments, Austin,
TX). The spectra were referenced to a standard flat surface of barium sulfate
Modeling Reflectance Spectra to Predict Discriminability
To construct a visually-perceivable model from fruit and background leaf
spectra, we estimated quantum catches by opsin Qi, where i is equal to each of the
535 nm, 550 nm, and 562 nm alleles present at the M/L locus in A. geoffroyi. For
A. palliatai, i is equal to 430 nm, 530 nm, and 560 nm. We followed Osorio
and Vorobyev [1996] in calculating fruit and leaf spectra:
Qi ¼
Ri ðlÞSðlÞIðlÞdl;
where l is the wavelength, and lmin and lmax are the lower and upper limits of the
visible spectrum for monkeys, respectively. Here we used lmin 5 390 nm and
lmax 5 700 nm. The spectral sensitivity of the ith opsin is Ri(l); the reflectance
spectrum of leaves or fruits is S(l), and I(l) is the illumination spectrum (i.e.,
source light). We assumed that the pre-retinal filtering was similar to that in the
human eye [Wyszecki & Stiles, 1982]. For A. geoffroyi we modeled the
performance of fruit detection from mature leaves depending on the opsin
present in the phenotypes.
Estimating the Performance of A. geoffroyi Phenotypes
We modeled the phenotype of the color vision following Riba-Hernández et al.
(this volume).
Calculation of Yellow-Blue and Red-Green Color Signals
Since color perception depends on the cone sensitivities present in the eye,
the classification of fruit by color is generally subjective. Based on quantum
catches by opsin calculations (QL, QM, and QS), color-vision channel information
can be estimated by
Yellow Blue Channel ðYBÞ ¼ QS=ðQL þ QMÞ
Red Green Channel ðRGÞ ¼ QL=ðQL þ QMÞ:
These two color-vision channels represent the neurological color-vision
path for the perception of color in primates [Regan et al., 2001]. The former
channel is present only in dichromatic individuals, while both channels
are present in trichromatic individuals. This analysis allowed us to define
fruit colors as either bluer or yellowier, and redder or greener than the leaf
background on a quantitative basis [Lucas et al., 2003; Regan et al., 1998].
We calculated the difference between fruit and background to define fruit
color. The difference between primates for each of the two color channels was
analyzed with the use of Student’s t-tests, and the data were log-transformed to
achieve normality. We used Systat 9.0 (SPSS Inc., Chicago, IL) for the
Am. J. Primatol. DOI 10.1002/ajp
404 / Stoner et al.
We collected a total of 900 hr of monkey feeding observations. Observation
times were approximately equal for both species (A. geoffroyi: 460 hr; A. palliata:
440 hr), and for both troops of A. palliata (222 and 218 hr, respectively). We
obtained only 60 hr of data on A. geoffroyi males, so we were unable to make any
comparisons between the sexes. Data were collected during 64 days of
observation. Ateles geoffroyi consumed a total of 63 fruit species, while A. palliata
consumed a total of 25 fruit species. For A. geoffroyi we collected 369 fruit
reflectance spectra from 25 species that represent 70% of the total fruit feeding
time. For A. palliata we collected 148 fruit reflectance spectra from 12 species
that also represented 70% of the total fruit feeding time. Fruit reflectances were
collected for all shared fruit species.
The two sympatric monkey species had very distinct diets. Alouatta palliata
consumed mostly young leaves, while A. geoffroyi consumed mostly fruits
(Table I). Alouatta palliata spent 34% of their total feeding time on fruits, while
60.5% of A. geoffroyi’s total feeding time was spent on fruits (Table I). These two
primates shared only seven fruit species, and these seven species were exploited
by them at different intensities (Morisita index 5 0.08; Table II). However, these
shared fruit species showed similar chromaticities (Table II, Fig. 1).
Contrary to our prediction, A. palliata did not consume more reddish fruits
than A. geoffroyi. Ateles geoffroyi primarily consumed fruits that were redder
than the background (62.2% of feeding time), while A. palliata dedicated only 24%
TABLE I. Comparative Diet of Ateles geoffroyi and Alouatta palliata Over an
Annual Period at Punta Rio Claro Wildlife Refuge, Osa Peninsula, Southwestern
Costa Rica
Plant item
Primate species
Mature leaves
Young leaves
Ateles geoffroyi
Alouatta palliata
The number of species is given in parentheses. Data based on 900 hr of observation.
TABLE II. Fruit Species Shared by A. geoffroyi and A. palliata and Their Average
Color7sd (red-green color channel)
Shared fruit species
Souroubea vallicola
Humiriastrum diguense
Byrsonima crispa
Licania sp. I
Spondias mombin
Dendropanax arboreus
Annual diet overlap (Morisita index)
A. palliata A. geoffroyi
Red-Green color channela
Values represent the percent of total foraging time (N 5 440 h for A. palliata and N 5 460 h for A. geoffroyi)
dedicated to each fruit species.
Background values range: 0.500–0.530.
Am. J. Primatol. DOI 10.1002/ajp
Use of Color Vision for Frugivory / 405
of their fruit feeding time to fruits of this hue (Fig. 1). In fact, the difference
between the fruit spectra and the background spectra (mature leaves) was
significantly greater for A. geoffroyi than for A. palliata (tstudent 5 –2.419,
P 5 0.021, d.f. 5 34.8). This difference was not significant for the YB color vision
channel (tstudent 5 0.809, P 5 0.430, d.f. 5 16). The most important fruits in the
diet of A. geoffroyi lay outside the chromaticities of mature leaves, while
A. palliata had more important species inside the background chromaticities
(Fig. 1). Ateles geoffroyi consumed a wider variety of fruit hues than A. palliata, as
can be seen by the greater distribution along the red-green axis of the food items
(Fig. 1).
Ateles geoffroyi
Allouata palliata
Fig. 1. Fruit chromaticities of the diet of A. geoffroyi (n 5 25 fruit species) and A. palliata (n 5 12
fruit species). The fruit chromaticities represent 70% of the total fruit diet for both monkey species,
and include the seven species that they shared. Circle size represents the percentage of each fruit
species in the diet of each primate (the bigger the circle the greater the percentage it represents).
Dashed square lines represent the chromaticities of a mature-leaf background. Fruit chromaticity
values 40.5 on the RG channel represent red fruit coloration. Circles with black dots represent
the fruit species that were shared by the two primate species (there are 14 black dots because
fruit chromaticities were measured for the seven shared species separately for A. geoffroyi and
A. palliata).
TABLE III. Fruit Detection Performance of Different Color Vision Phenotypes of
A. geoffroy
All Trichromats
Dichromat 562
Dichromat 535
Dichromat 550
Percent of fruit species distinguished
Numbers represent the percentage of fruit species (N 5 39) in which the mean signal (jnd values) exceeded a
value greater than 1 jnd.
Am. J. Primatol. DOI 10.1002/ajp
406 / Stoner et al.
The model predicted that trichromat phenotypes would detect the total
number of fruit species used by A. geoffroyi, while dichromats would detect 95%
of the fruits in their diet. Nevertheless, trichromats outperformed dichromats
for 23 species (Table III), while dichromats performed equally to trichromats for
14 species. In any case, trichromats outperformed dichromats for fruit detection.
The reason for the maintenance of opsin gene allele polymorphism remains
unclear [Cropp et al., 2002; Dominy et al., 2003; Surridge et al., 2003]. According
to the historical hypothesis, frugivorous species would be expected to be routine
trichromats. Paradoxically, in platyrrhines, most frugivorous species are polymorphic, while the only routine trichromat, Alouatta spp., is mostly folivorous. In
fact, it has been suggested that platyrrhine polymorphism is inadequate for a
frugivorous diet [Regan et al., 1998]. On the contrary, we found that
polymorphism in platyrrhines appears to be well suited to the task of fruit
detection. In fact, dichromatic phenotypes can detect almost all of the species in
the diet of A. geoffroyi, but trichromats are better at this task [Riba-Hernández
et al., 2004]. Our results support the hypothesis that trichromacy is advantageous
for frugivory in platyrrhine monkeys, with the corollary that the performance of
dichromats does not appear to be as poor as has been suggested. As predicted,
A. geoffroyi consumed fruits with a wider variety of hues than A. palliata.
Nevertheless, it is not clear whether this is because dichromatic individuals that
cannot distinguish red hues randomly consume a greater variety of colors because
they do not all have the capacity to distinguish the critical hues, or simply because
Ateles consumes more fruit (not being able to fall back on leaves) and
consequently a greater number of species (63 vs. 25 for Alouatta).
The advantage of trichromacy over dichromacy in platyrrhines may lie not
only in the discrimination performance, but also in the information that the
discrimination provides about the quality of the resource being exploited
[Dominy, 2003; Dominy & Lucas, 2001, 2004] (Riba-Hernández et al., this
volume). For A. geoffroyi, glucose-rich fruit species are better detected by
trichromats than dichromats (Riba-Hernández et al., this volume); therefore,
trichromats may lead dichromats to patches of glucose-rich fruits [Lucas et al.,
2005]. This foraging pattern may be sufficient to maintain the M/L polymorphism
of platyrrhines by a heterozygous advantage [Mollon et al., 1984], if feeding on
fruits is the selective pressure behind the evolution of color vision in primates.
This pattern would also explain the fact that redder fruits were more common in
the diet of A. geoffroyi.
One possible explanation for the fact that trichromatic A. palliata did not
consume more reddish fruits involves physiological constraints. Alouatta spp. are
hindgut fermenters, and this kind of digestion system is severely affected by high
sugar intake, which limits the effectiveness of hindgut fermentation [Edwards &
Ullrey, 1999; Milton, 1998]. Although A. palliata shared fruit species with
A. geoffroyi (Table II), these species are among the species that show lower values
of sugar concentration and a greener/yellowier coloration (rather than red) [RibaHernández et al., 2004]. Furthermore, it has been shown that the red-green color
channel is positively correlated with glucose concentration in fruits consumed by
A. geoffroyi at our study site (Riba-Hernández et al., this volume). If hindgut
fermentation limits the amount of sugary fruits that A. palliata can consume, it is
possible that they are actually using their trichromatic ability to discriminate
Am. J. Primatol. DOI 10.1002/ajp
Use of Color Vision for Frugivory / 407
reddish colored fruits to avoid consuming fruits of this hue and the subsequent
high concentrations of glucose.
Another explanation for the finding that A. palliata did not consume more
reddish fruits at our study site may lie in competitive interactions with
A. geoffroyi. Alouatta spp. are usually displaced by other primate species from
feeding sites (Stoner, unpublished data) [Stevenson et al., 2000], which reduces
the overlap of diet between these two primates. This suggests that trichromacy
may be more relevant to Alouatta spp. for folivory than for frugivory, by enabling
them to identify young leaves that are often more reddish than mature leaves
[Stoner et al., 2000]. At our study site, Stoner et al. [2000] found that the selection
of young leaves by A. palliata was highly related to the nutritious quality of the
leaves, especially in terms of proteins and lower concentrations of phenols,
similarly to patterns reported for catarrhine monkeys [Dominy & Lucas, 2001].
Comparative studies of sympatric species have provided new insights into the
evolution of color vision in primates [Dominy & Lucas, 2001; Smith et al., 2003;
Sumner & Mollon, 2000a,b]. Here we present new information on the use of
trichromatic color vision by primates. We suggest that some species of primates,
such as Alouatta spp., may actually use their visual discrimination to avoid
reddish fruits. We further suggest that other factors, such as competitive
interactions among species, may play important roles in shaping the actual
patterns of color vision in extant primates. More detailed studies on sympatric
platyrrhine species at other sites are needed to identify the importance of
competitive interactions between primate species as a selective pressure in
shaping the evolution of color vision in primates.
We thank Hannah Buchanan-Smith and Valdir F. Pessoa for the invitation to
participate in the ‘‘Recent Advances in Color Vision Research’’ symposium held
at the 20th Annual Congress of the International Primatological Society, 2004.
We thank Silvia Solis Madrigal and Wanda Petersen Pereira for their invaluable
field assistance. We also thank Mauricio Jimenez for his veterinary support in
tranquilizing and placing radiocollars on the howler monkeys, Daniel Osorio for
his help with the color modeling, and the Ministerio del Ambiente y Energı́a for
permission to conduct this research. We are grateful to two anonymous reviewers
for helpful comments that improved the manuscript, and H. Ferreira and
G. Sánchez Montyo for technical assistance.
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Am. J. Primatol. DOI 10.1002/ajp
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species, color, sympatric, vision, platyrrhine, comparative, use, frugivores
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