Geographic Range
There are three recognized subspecies of mustached tamarins:
Saguinus mystax mystax
,
Saguinus mystax pileatus
, and
Saguinus mystax pluto
. All three subspecies are found in the middle Amazon region of northern Peru and
western Brazil.
- Biogeographic Regions
- neotropical
Habitat
Mustached tamarins prefer dry, upland forests in the Amazonian lowland and avoid flooded
forests.
- Habitat Regions
- tropical
- terrestrial
- Terrestrial Biomes
- rainforest
Physical Description
Mustached tamarins, being
callitrichines
, have distinguishing features that set them apart from other
New World monkeys
. These include their diminutive body size and the presence of claw-like nails on
all digits except the hallux. Mustached tamarins have also lost both their upper and
lower 3rd molars and the hypocones on their upper molars. The dental formula is 2.1.3.2
/ 2.1.3.2. Tamarins have spatulate incisors that are shorter than their canines, unlike
marmosets
which have incisors and canines of more similar size. The labial and lingual sides
of their incisors are also thickened with enamel. Mustached tamarins have jet black
pelage on their body and tail and a characteristic white stripe on their upper lip,
which gives the appearance of a "white mustache." Mustached tamarins do not have prehensile
tails.
Mustached tamarins do not exhibit sexual dimorphism and there are only minor differences
in body and canine size between males and females. On average, adult male mustached
tamarins weigh 491 ± 23.0 g and measure 61.2 ± 31.2 cm (n=95), whereas adult females
weigh 511.0 ± 49.0 g and measure 62.0 ± 29.0 cm (n=80).
According to Cesar and Bicca-Marques (1999), feeding behaviors are correlated with
the relative size and shape of hands in
callitrichines
. Mixed-species groups of tamarins are observed to form only between species that
have different foraging behavior, which reduces direct competition. Consequently,
due to the specialization conferred by hand morphology, species with dissimilar hand
shapes are usually found in association with each other in such mixed-species groups.
This is the case in the mixed-species groups of mustached tamarins and saddle-back
tamarins.
Saguinus mystax
captures exposed insects on branches and thus has evolved relatively wide and short
hands, as opposed to
saddle-back tamarins
, which have made use of longer and more slender appendages to manipulate openings
in tree trunks and other substrates to search for insect prey.
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes alike
Reproduction
Mustached tamarins usually associate in stable troops, which vary widely in size.
For example, the mean group size was found to be 6 to 7 individuals per group in two
studies (Garber et al., 1993; Löttker et al., 2004), with a range of 4 to 11 individuals
in each group.
The reproductive strategy of
S. mystax
is described as polyandrous or polygynous. Each group is typically centered on one
breeding female, which is usually also the oldest female. Despite the lack of breeding
opportunities, adult non-breeding females may continue to persist in the group, and
may ascend to breeding female status in the event of death or illness of the current
breeding female.
A study by Löttker et al (2007) suggests that breeding females use social grooming
as incentive to induce adult males and non-breeding offspring to stay in the group
as helpers to care for their young.
Adult mustached tamarins live in highly tolerant and cooperative societies. Garber
(1997) estimated that the frequency of cooperative to aggressive acts between adult
mustached tamarins was 52:1. Intragroup sexual aggression between male mustached tamarins
is exceedingly rare, but mate guarding may be practiced by the dominant breeding adult
male during the breeding female’s most fertile periods.
Male mustached tamarins may compete for reproductive success via sperm competition
and other physiological mechanisms. For example, testes volume in males within a group
varied greatly among individuals and this could directly impact reproductive viability
of males.
Despite a polyandrous mating system and seemingly a lack of evidence of strong competition
or aggression over breeding opportunities, a genetic analysis of paternity in mustached
tamarins revealed that 67 to 100% of
S. mystax
infants in a troop were fathered by the same adult male. Genetic relatedness within
a troop is high but both non-related and related adults can co-exist in a troop. There
is a need for a better understanding of the critical factors that govern the observed
reproductive bias among adult
S. mystax
males.
- Mating System
- polyandrous
- polygynous
- polygynandrous (promiscuous)
- cooperative breeder
Mustached tamarins have characteristic twinned births, with an annual birth peak from
November through March. Some
S. mystax
troops experience 2 birthing cycles, from June to October and in February and March.
Spatio-genetic analysis of a population of mustached tamarins in Peru by Huck et al.
(2007) revealed variable genetic relatedness within a troop. Migration is common among
male and female mustached tamarins, but females are often required to travel for longer
distances than males before settling down with a new troop due to scarce breeding
opportunities in a polyandrous mating society. Matings are generally between non-closely
related partners, hence the degree of inbreeding is reduced.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- viviparous
Maternal infanticide, while rare, has been documented twice in Saguinus species, in S. mystax by Culot et al. (2011) and S. fuscicollis by Herrera et al. (2000). In both cases, infanticides occurred under circumstances of low availability of helpers (2 to 3 male helpers compared to 4 to 5 helpers at other times) and presence of more than 1 breeding female in the troop. Both of these factors lower the likelihood of obtaining sufficient care for the infant within the group.
The single most important determinant of maternal infanticide could be the mother’s
perception of low probability of survival of her infant in the group, which is directly
caused by external factors in the troops, such as low helper availability. Low helper
availability or competition for care with infants from another breeding female in
the same troop greatly reduces the chances of survival for infant mustached tamarins.
Maternal infanticide may thus represent a strategy to maximize efforts that go into
ensuring survival of infants. On the contrary, infanticide of unrelated infant tamarins
by male mustached tamarins, which is usually attributed to sexual selection pressures,
has never been observed in
Saguinus
. Males do not gain reproductive advantages from infanticide, as infant mortality
has minimum effect on the mother’s fertility status. Female mustached tamarins remain
reproductively viable during nursing of their young.
Due to the cooperative breeding behavior of
S. mystax
, the number of helpers available in a group to care for the young has a strong influence
on the survival of infant tamarins (Garber, 1997).
Birth intervals were also shorter during the periods when infanticide occurred (1
to 2.5 months). In
Saguinus fuscicollis
, which also practices cooperative breeding like
S. mystax
, the optimal birth spacing is 3 months, so as to allow for efficient care of the
young. Shorter birth intervals increase the burden on the troop to care for the young,
and this effectively reduces the likelihood of providing adequate care for all of
them.
Although not demonstrated in
S. mystax
, care and transport of infants results in significant personal costs for small-sized
mustached tamarins. Loss of body mass of carriers of infants has been demonstrated
in wild and captive
Saguinus oedipus
. Cooperative breeding allows the costs of caring for the young to be distributed
to other members of a troop to a certain extent, and carriage of infant mustached
tamarins is not restricted to certain members of a troop, as members usually take
turns in transporting the young when foraging.
- Parental Investment
- precocial
- male parental care
- female parental care
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- male
- female
-
protecting
- male
- female
-
provisioning
- post-independence association with parents
Lifespan/Longevity
There are inconclusive data on the longest lifespan of mustached tamarins in the wild,
though individuals in captivity have been known to live in excess of 20 years.
Behavior
Mustached tamarins are social and diurnal animals and usually co-exist in stable groups
characterized by infrequent conflicts and relatively low occurrence of sexual interactions
within the group.
Being arboreal, mustached tamarins usually occupy the middle and upper layers of the
forest at a height of 10 m above the ground. This is in contrast to their congener,
saddle-back tamarins
, which occupy the lower layers of the forest.
Aggressive intergroup encounters between mustached tamarin groups result primarily
from resource and mate defense and less from territorial disputes.
Interestingly, it was observed that in mixed-species groups, the larger
S. mystax
members are more likely to gain access to disputed food resources over
Saguinus fuscicollis
members in the same troop.
Conflicts between neighboring troops of mustached tamarins frequently occur over food
resources and especially near important feeding trees. The loss of such feeding opportunities
will result in extra foraging efforts by the displaced troop. According to Garber
(1988b, p. 27-28), "in those instances in which the troop was excluded from using
these nearby feeding sites, mean distance traveled to the next major feeding tree
was 331 m. This is considerably greater than the distance traveled to the next major
feeding site (59.2 m) after a successful defense." Hence, mustached tamarins that
are displaced from feeding by another troop will have to expend more energy to travel
longer distances to find another suitable feeding site.
The development of trichromatic vision in primates has been proposed as an adaptation
to allow better detection and identification of fruits in their forested habitats.
New World monkeys
, such as
Saguinus
, are unique in having a polymorphic vision system. Only heterozygous female mustached
tamarins have trichromatic vision, with the rest retaining dichromatic vision. There
is some evidence in the closely related species
Saguinus labiatus
that those with trichomatic vision "were more efficient at selecting ripe fruits
than were dichromats" (Smith et al 2003, p.3159), but this has not been independently
demonstrated in
S. mystax
.
According to a study by Smith et al. (2003b), mixed-species troops of mustached and
saddle-back tamarins were usually led by a mustached tamarin during migration through
its home range. Color vision status did not consistently influence the choice of the
lead animal of a troop. However, it was observed that a trichromatic female was significantly
more likely to lead when the troop was approaching a fruiting tree. This lends support
to the hypothesis that trichromats are more endowed to differentiate between fruiting
and non-fruiting trees, and thus would be able to identify fruiting trees that are
ready for feeding. Color vision is likely to be just one of many factors affecting
leadership selection in wild
S. mystax
and
S. fuscicollis
troops.
Vigilance, defined as "observable head movements in stationary animals where scanning
is directed beyond arm's reach" (Stojan-Dolar and Heymann, 2010b, p. 327), appears
to serve the primary function of predator detection and avoidance in
S. mystax
. Secondarily, vigilance has also been proposed to maintain group cohesion within
one’s own group and to monitor the movement of neighboring troops. Group composition
and size affect the vigilance of mustached tamarins. Individuals in mixed-species
groups appeared to be less vigilant when feeding and during periods of resting than
those not in mixed-species groups.
Saguinus mystax
members in mixed-species groups also exhibited a greater reduction in vertical vigilance,
compared to that in the horizontal direction. One reason may be the positional preferences
of
Saguinus mystax
and
S. fuscicollis
, which occupy different forest strata. Tamarins in mixed-species groups may share
responsibility for group vigilance in the vertical direction; hence, greater reduction
in vertical scanning is observed. Development of mixed-species groups provides some
evolutionary advantage in enhancing predator detection and avoidance for the troop
so as to increase the chances of overall survival of its
S. mystax
and
S. fuscicollis
members.
Symmetrical quadrupedalism is the dominant type of locomotion in
S. mystax
(63%), followed by asymmetric quadrupedalism (23%) and leaping (14%). Mustached tamarins
also preferentially use more horizontal supports when traveling, and thin and flexible
supporting structures when feeding.
The movement of
S. mystax
has been described by Sussman and Kinzey (1984, p. 427) as “quadrupedal, running,
and leaping along medium to small branches and jumping among the fine terminal branches
and leaves between trees.”
Although other primates do form sympatric groups, the mixed-species groups of tamarins
are among the most stable and permanent of all primate groups. Group cohesion is important
in maintaining mixed-species groups, and this can be enhanced by intra-group communication
and shared responsibility in predator defense and avoidance. As tamarin species in
mixed-species groups may compete for similar resources, spatial segregation between
species is important in allowing such mixed-species groups to persist.
By differing in locomotor types, support preference, and food/prey selection, and
by occupying different strata of their forest habitat, mustached tamarins and
saddle-back tamarins
are able to co-exist harmoniously in groups without competing for the same scarce
resources.
Home Range
The home range of mustached tamarins is highly variable, and several troops may co-exist
with overlapping territorial limits. A study of mustached tamarins inhabiting Padre
Isla (in the Amazon Basin) showed that home ranges of neighboring
S. mystax
groups overlapped by approximately 38%.
Communication and Perception
Allogrooming and scent marking have been studied in
S. mystax
societies. Allogrooming or social grooming is commonly observed in mustached tamarins,
but it is not practiced equally by all members of a troop. In one study by Heymann
(1996), an adult male contributed 70% of all grooming observed in a group, without
bias to any one of the other members in his troop. The reasons for this asymmetry
are not well-characterized but may be attributed to differences in social standing
or competition for reproductive success. As mentioned above, breeding females may
employ social grooming as a way of encouraging other members in her troop to stay
to assist in parental care duties.
The use of scent marking for territorial demarcation is variable among
Saguinus
species. In both wild and captive
S. mystax
, there is little evidence for the use of scent marking in territorial defense.
Scent marking functions prominently in intra-group communication and female mustached
tamarins are more likely to engage in scent marking behavior, compared to males. On
the contrary, male mustached tamarins are more likely to respond to scent marks of
the opposite sex, which may be related to mate selection. Mustached tamarins do not
engage in "collective scent marking" or allomarking, unlike in
S. fuscicollis
.
The rates of scent marking by mustached tamarins are directly proportional to the
duration of occupation of areas by the troop. Intense scent marking in more frequented
areas enhances the effectiveness of olfactory communication between members of the
same troop.
Vocalization has also been demonstrated to be a form of communication in
S. mystax
, such as in situations where a mustached tamarin becomes separated from its troop.
According to Snowdon and Hodun (1985, p. 211), "The long-call structure of the
moustached tamarin is similar to that of its sympatric tamarin species,
saddleback tamarins
. In both species the calls last 2 to 3 seconds, consist of several short, frequency-modulated
syllables, and has a frequency range of 8 to 12 kHz". The same study also noted
that "nonbreeding adult females gave fewer calls" and they were also slower to make
the first long call upon separation. This could reflect "the relatively low status
of nonreproductive mature females" (p. 211) in their troop, which is usually centered
on a single breeding female. Members of a troop are able to recognize members of their
troops by responding to long calls from separated members. This ability to distinguish
individual members suggests that a
S. mystax
troop in the wild is able to maintain its integrity by ensuring that all members
are accounted for.
Early morning vocalizations and subsequent coordinated movement patterns within mixed-species
troops of mustached tamarins and saddle-back tamarins are also crucial in maintaining
group integrity on a daily basis.
- Other Communication Modes
- scent marks
Food Habits
Being frugivorous-insectivorous, mustached tamarins feed on fruits, flowers, and nectar,
insects, and plant exudates. Food sources like gums and soil appear to supplement
nutrients and minerals lacking in their main diet of fruits and insects.
Mustached tamarins typically only feed on a small number of plant species in their
home range. However, feeding trees are numerous and scattered throughout their home
range. Feeding trees are usually continuously fruiting and produce a small amount
of fruits daily, hence mustached tamarins may visit the same trees regularly over
a span of several months to exploit the resources.
In a group of mustached tamarins in northeastern Peru, up to a third of feeding time
was spent feeding on nectar from
Symphonia globulifera
in the dry season. Mustached tamarins concentrated their foraging efforts on trees
that yielded more nectar, despite having to travel greater distances between feeding
sessions in some cases. Therefore, proximity of feeding trees from one another did
not predict foraging patterns, as mustached tamarins preferentially chose feeding
trees that allowed them to feed for longer periods of time instead of trees that were
closer and would thus minimize traveling time from one tree to another.
Although feeding trees are typically scattered throughout their home range, mustached
tamarins are able to remember the locations of the important feeding trees to minimize
efforts in foraging. The use of spatial memory and learning in foraging are important
because they provide "an efficient means of exploiting a widely scattered set of predictable
feeding sites" (Garber, 1989, p. 212).
Gum feeding is highly seasonal, but it is a major dietary staple during dry and early
wet seasons when other food sources are scarce. Furthermore, gum exudates balance
mineral intake, which will otherwise suffer from the effects of a low-calcium and
high-phosphorus insectivorous diet.
Since mustached tamarins lack the dental specialization to gouge holes into tree bark,
gum feeding is "dependent principally on either natural weathering of the bark or
on the parasitic activity of woodboring insects, or both" (Garber, 1992, p. 471).
Heymann and Smith (1999) found that
Saguinus mystax
concentrated gum feeding in the afternoon. They hypothesized that this may be "a
behavioral strategy to prolong the time that the gums are retained in the gastrointestinal
tract" (p. 468). Prolonged passage allows for better digestion and more complete microbial
fermentation of the substrates. This in turn yields nutritional benefits while providing
these small animals with a means to generate heat during the night via the fermentation
process.
Garber (1992) described mustached tamarins as exhibiting "a highly opportunistic foraging
pattern," with "seasonal exudate feeding, occasional trunk foraging patterns, and
small-branch fruit and insect feeding" (p. 471).
The main vertebrate prey for mustached tamarins includes "katydids (
Tettigoniidae
,
Orthoptera
), stick grasshoppers (
Proscopiidae
,
Orthoptera
), and spiders (
Araneidae
), although lizards (
Squamata
) and frogs (
Anura
) were also taken" (Smith, 2000, p. 315). Mustached tamarins also feed on nestling
birds, albeit less frequently. They forage for prey at higher levels within the forest
strata than
Saguinus fuscicollis
(average of 8.84 m vs. 1.50 m; Smith 2000), and also capture most prey from slender
and horizontal substrates.
- Primary Diet
- omnivore
- Animal Foods
- birds
- amphibians
- reptiles
- insects
- Plant Foods
- fruit
- nectar
- flowers
- sap or other plant fluids
Predation
Due to their small size, mustached tamarins are vulnerable to predation by a large
number of terrestrial predators, such as tayras (
Eira barbara
), ocelots (
Leopardus pardalis
), and jaguarundis (
Puma yagouaroundi
). Large snakes like
Boa constrictor
and birds of prey are also important predators.
Cooperative defense against predators was observed in a tribe of mustached tamarins
in northeastern Peru. Members of this tribe successfully rescued a mustached tamarin
that was being attacked by a
Boa constrictor
by counter-attacking the predator. The tribe subsequently avoided low heights for
a week after the attack, as further deterrence to attacks by snakes.
In general, mustached tamarins counter predators by avoiding attracting attention.
Mustached tamarins are camouflaged in densely foliated areas and they carefully select
resting and sleeping sites that have lower chances of being exposed. They may also
choose resting sites that are physically isolated from surrounding vegetation, as
this would increase the likelihood of them being able to detect approaching predators.
- Anti-predator Adaptations
- aposematic
- cryptic
Ecosystem Roles
Mustached tamarins play an important ecological role in their habitat by dispersing
seeds of plant species in primary and secondary rain forest, and thus indirectly impact
forest regeneration and maintenance.
As mustached tamarins are opportunistic feeders and are able to explore many kinds
of plant resources, they are able to disperse a great variety of seed species.
Due to their small size, there is a limit to the size of seeds that they can disperse.
Seasonal changes in diet, sleeping and resting patterns, and variation in foraging
behavior and migration may also affect their contribution to seed dispersal in the
rainforest.
On a negative note, mustached tamarins have been observed to be highly destructive
in their nectar-feeding activities but the widespread ecological impact of this has
not been demonstrated.
Mustached tamarins that foraged near human settlements were found to have higher prevalence
of a primate parasite,
Prosthenorchis elegans
. However, in the same study by Wentz et al. (2010), mustached tamarins were not found
to be infected by human parasites even though they lived in close proximity to human
communities with high rates of infestation of nematodes such as
Ascaris lumbricoides
and hookworms.
- Ecosystem Impact
- disperses seeds
Economic Importance for Humans: Positive
Saguinus mystax
, like other tamarin species, has been used extensively in biomedical research, e.g.,
in the development of the hepatitis A vaccine.
- Positive Impacts
- research and education
Economic Importance for Humans: Negative
Although commensally associated with human communities, there is no evidence for adverse economic effects on humans.
Conservation Status
Mustached tamarins are on the least concern list of animals on The IUCN Red List of
Threatened Species (last assessed in 2008). Populations of wild mustached tamarins
have been noted to be stable, although habitat destruction remains a threat to species
living in the Amazonian rainforest.
The effects of agriculture on
S. mystax
populations in Peru have been studied. Populations recovered and even exceeded pre-agricultural
numbers due to "increased reproductive rate, early breeding, and reduced infant mortality",
while "immigration seemed to play a minor role" (Ramirez, 1984, p. 258).
Other Comments
Genetic analysis supports the division of tamarin taxa into small-bodied and large-bodied
clades (Matauschek et al., 2011, p. 564), of which
S. mystax
is a member of the large-bodied clade. Molecular phylogenetic evidence also supports
the “phyletic dwarfism” hypothesis in
Callitrichinae
, which is "defined as a gradient in morphological size partially correlated with
evolutionary time" (Perelman et al., 2011, p. 4). In this case,
Saguinus
is the first to diverge, followed by the remaining
callitrichines
in order of decreasing body size. Within
Saguinus
, phylogenetic analysis of mitochrondrial DNA has demonstrated that
Saguinus labiatus
is the closest relative of
S. mystax
, with the two taxa diverging about 1.15 mya.
Additional Links
Contributors
Jayne Lim (author), Yale University, Eric Sargis (editor), Yale University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Neotropical
-
living in the southern part of the New World. In other words, Central and South America.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- polyandrous
-
Referring to a mating system in which a female mates with several males during one breeding season (compare polygynous).
- polygynous
-
having more than one female as a mate at one time
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- cooperative breeder
-
helpers provide assistance in raising young that are not their own
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- young precocial
-
young are relatively well-developed when born
- male parental care
-
parental care is carried out by males
- female parental care
-
parental care is carried out by females
- arboreal
-
Referring to an animal that lives in trees; tree-climbing.
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- scent marks
-
communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- aposematic
-
having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- omnivore
-
an animal that mainly eats all kinds of things, including plants and animals
References
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Buchanan-Smith, H., S. Hardie, C. Careres, M. Prescott. 2000. Distribution and forest utilization of Saguinus and other primates of the Pando Department, Northern Bolivia. International Journal of Primatology , 21/3: 353-379.
Buchanan-Smith, H. 1990. Polyspecific association of two tamarin species, Saguinus labiatus and Saguinus fuscicollis , in Bolivia. American Journal of Primatology , 22/3: 205-214.
Cesar, J., J. Bicca-Marques. 1999. Hand specialization, sympatry, and mixed-species associations in callitrichines. Journal of Human Evolution , 36/4: 349-378.
Culot, L., Y. Lledo-Ferrer, O. Hoelscher, F. Munoz-Lazo, M. Huynen, E. Heymann. 2011. Reproductive failure, possible maternal infanticide, and cannibalism in wild moustached tamarins, Saguinus mystax . Primates , 52/2: 179-186.
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