Nocturnality in the Owl Monkey, Aotus sp.

Jonathan Greenberg
3 December 1997

Introduction
Aotus sp., the owl monkey, is the worlds only nocturnal monkey and one of the most
widespread primates in the neotropics. The odd activity pattern of this primate seems at first
anomalous but at a closer glance brings up many proximate and ultimate level questions as to its
development and yields insights into wider-ranging ecological principles. For a primate which
shares similar morphology, social system and diet to other diurnal primates such as the titi monkey
(Callicebus), Aotus has markedly different lifestyle strategies. Morphology, geographical and
habitat distribution, social system and diet all factor into the explanations of the development of
nocturnality on Aotus. These differences can serve to test the effects of interspecific competition,
predation, group size and foraging patterns.

A strong case can be made for the effects of

interspecific competition being the driving force behind Aotus activity patterns by examining
variation both intraspecifically as well as between Aotus and other sympatric primates. Dietary
choices are also suggested to play an important role in determining activity patterns. Predation
pressure differences between night and day may also causes an adaptive response leading towards
activity patterns matching periods of low predation.

Morphology
Aotus sp. is a relatively small-bodied primate. Head and body measures approximately
350 mm, weight 1 kg. (Napier & Napier, 1996). Sexual dimorphism is almost nonexistent. The
coats are thick and counter shaded: grayish brown with pale underparts (Napier & Napier, 1996).
Aotus coats are relatively cryptic and resemble many other nocturnal animals including lorises,
phalangerids, opossums and owls (Moynihan, 1976). The faces have dark stripes and white
patches over the eyes. The white patches may be used in intraspecific encounters or for defense to
give the illusion of looking alert even while their eyes are closed (Moynihan, 1976). The eyes are

large and lack cones but contain a fovea, suggesting the ancestor of Aotus was a diurnal monkey
(Napier & Napier, 1996). Aotus is color blind. The tail is nearly as long as the body and acts as a
balancing organ for quadrupedal locomotion (Wright, 1985). Aotus intermembranal index is 74
(Napier & Napier, 1996). The ears are small and round (Wright, 1985). The digits of Aotus have
true nails, two of which are possibly a grooming nail on the toes similar to those found in
prosimians (Wright, 1985). Aotus has conspicuous tactile pads (Wright, 1985), the terminus of
which is free to bring into contact with branches and other objects in its environment (Napier,
1976). Teeth are adapted to frugivory: canines are small and ineffective as nut-crackers and molars
have a relatively unpronounced cristid obliqua (Wright, 1985). The mandible is more shallow than
Alouatta and Callicebus, suggesting little specialization to folivory (Wright, 1985).

The

vomeronasal organ and nasopalatine ducts of Aotus are large (Hunter et al., 1984) and olfaction is
well-developed when compared to other Neotropical primates (Bolen and Green, 1997).

Taxonomy
Aotus sp. (Family Cebidae, Subfamily Aotinae) is most closely related to the titi monkeys
(Callicebus sp.). Originally thought to be a single species with different subspecies, more recent
analysis has split the genus into at least nine distinct species (Hershkovitz, 1983). The genus
Aotus may be over 12 million years old (Setoguchi and Rosenberger, 1987) and probably evolved
from a diurnal ancestor into nocturnality which was, at the time, practiced solely by marsupials and
rodents (Wright, 1995), most of which are quite a bit smaller than Aotus.

Setoguchi and

Rosenberger (1987) suggest Aotus nocturnal lifestyle dates back at least as far as the 12 million
year old fossil of Aotus didensis.

Geographical Distribution and Habitat

Aotus is found in most of northern South America from Panama to northern Argentina and
from the Pacific in Ecuador and Colombia to the Atlantic in Brazil (Robinson et al., 1987). They
are found from lowland forest to high altitude cloud forests at 3000 m (Napier, 1976). They can

occupy virtually every wooded habitat found in these regions. Aotus are found in every strata of a
forest, from 7 to 35 m (Wright, 1981).
Aotus sleeps in holes in hollow trees or in dense vine tangles (Wright, 1981). There is
some variance in nesting preference geographically and Rathbun and Gache (1980) have reported
Argentinean owl monkeys sleeping almost exclusively in vine tangles. There are reported cases of
interspecific competition over sleeping sites with Potos flavus (kinkajou) (Aquino and
Encarnacion, 1986). Cohabitation with other species has also been reported with bats, P. flavus,
Bassaricyon gabbi (olingo), Coendu bicolor (Aquino and Encarnacion, 1986), and Isothrix
bistriatus (Puertas et al., 1995). Coocupation of the same tree has been seen with Coendou sp.
and Caluromys lanatus (Puertas et al., 1995). Choice of concealed nesting sites is probably
influenced by predation (Heymann, 1995 and Isbell, 1994) and proximity to feeding sites
(Heymann, 1995).
Aotus is found in regions with extremes in temperatures, particularly in the subtropical dry
forests of the Chaco of Paraguay where daily temperatures can range from 40 C down to -5C
(Wright, 1995).

Social System
Aotus is usually reported as a highly monogamous primate. Groups are often composed of
an adult male and female and up to three infants and juveniles. However, reliable observations of
up to five multi-adult groups moving through the canopy together have been reported (L. T.
Rosengreen, pers. comm.) and multiple-adult groups nesting together (Hernandez-Camacho and
Cooper, 1976). Other reports of larger groups of up to 30 individuals found in fruiting trees are
probably aggregations of smaller groups, a point supported by a high degree of agonistic behavior
between individuals in the larger groupings (Wright, 1981).
Owl monkeys have a relatively short interbirth interval of about one year which is possible
due to high levels of paternal care which alleviates much of the energetic costs to the mother
(Garber and Leigh, 1997). Male owl monkeys carry the young, defend them from predators, play

and instruct them (Wright, 1985). The social system benefits the male, usually limited by access to
females, by providing high parental certainty (Garber and Leigh, 1997), low infanticide rates and
short interbirth intervals. Owl monkeys are unique in that females will actively refuse to carry the
young if the male is unavailable, going so far as to violently pull the infant off of its back (Eduardo
???, pers. comm.) Two birth peaks occur, one at the end of the dry season and the other in the
middle of the wet season (Wright, 1985). Gestation is about 133 days (Hunter et al., 1979).
Young disperse at two to three years (Wright, 1985) and probably pass into a nomadic,
vagabond stage before pair-bonding with a female (Charles-Dominique, 1977).
Male-male aggression is common and is a factor in keeping groups apart (Moynihan,
1964). Agonistic encounters involve back arching, stiff-legged jumping, pilo-erection, urination
and defecation, as well as giving clicking/grunting alarm calls. Identical agonistic displays occur
between conspecifics and other species (Wright, 1978).
Aotus is highly territorial. Territories are extremely small for a primate of its size, usually
no greater than 10 ha (Wright, 1985). Although little work has been done on the dynamics of
territory size and location over time, it has been suggested, based on the behaviors of nocturnal
prosimians, that Aotus is relatively sedentary (Charles-Dominique, 1977). This is supported by
Isbells (1994) conclusion that use of unfamiliar areas by primates may increase the rates of
predation. Relatively mobile, daily path lengths are reported to be approximately three-quarters of
a kilometer. Territories between neighboring groups overlap extensively (Wright, 1978). Aotus
has been reported to come out of trees and cross open savanna to move between forest patches
(Rathbun and Gache, 1977). Population densities in Peru are reported to be around 25 to 50
individuals per km2 (Moynihan, 1976).
Aotus is a relatively noisy monkey, uttering loud contact and locomotory notes (Moynihan,
1976). Olfaction is an important component of communication and Aotus marks substrates by
rubbing a gland at the base of its tail and exuding a brown, oily substance (Wright, 1981).

Diet

Aotus diet consists of fruits, insects and leaves. Spending approximately 50% of its
waking hours foraging (Wright, 1985), the owl monkey has a wide range of diet compositions
both geographically as well as seasonally.

In Peru, Aotus spends three-quarters of its time

foraging for fruits, and the remaining time searching for leaves and insects.

In Paraguay,

conversely, only 16% of its time is spent foraging for fruits, 40% on leaves, 11% on insects and
the remaining 33% on other food sources (Wright, 1985).

Aotus spend much of their time

foraging in medium- to large-crowned trees.

Fruits are usually sweet or bland, though about 15% are bitter-tasting . There is probably
an increased ability to digest secondary compounds found in the fruits and other foods consumed.
Due to the color-blindness, color choices are presumed to be irrelevant. Aotus feeds on over 70
different species of fruits in Peru. Figs become extremely important food sources in time of
scarcity (Wright, 1985).
Due to the small body size, folivory is uncommon in wetter habitats. Leaves eaten are
either young leaves or taken from lianas of any age. Flowers and nectar are eaten when available,
and nectar becomes the sole source of nourishment in July and August when food is extremely
scarce (Wright, 1985). In areas with little fruits or nectar in scarce seasons, folivory increases
significantly (Wright, 1995).
Seed predation is rare in Aotus. The seeds pass unharmed through the gut (Wright, 1985)
which may suggest Aotus is an important dispersal agent.
Insects are an important source of protein for the owl monkey and occupies, on the
average, one-fifth of their foraging time. Over half of the feces collected for Aotus contained
insect parts, compared to less than one-fifth for Callicebus (Wright, 1985). Aotus preys primarily
on large orthopterans, lepidopterans, coleopterans and spiders which it forages for at dusk and
dawn and on clear, moonlit nights (Wright, 1995). Rarely missing, Aotus captures insects during
methodical movements along branches and catching them on the branches or in the air (Wright,
1985).

There have been reports of consumption of vertebrates such as bats, small birds, eggs and
lizards (Bates, 1863; Sanderson, 1957) but this is probably rare (Wright, 1985).

Predation
Predation pressures are suggested to be low for Aotus.

Nocturnal activity patterns

preclude many of the diurnal predators, such as raptors, from creating any serious threat.
Nocturnal prey, also, are harder to detect than are diurnal (Isbell, 1994). Although occasionally
nocturnal, many of the potentially threatening snakes are terrestrial with few exceptions, and Aotus
rarely leaves the trees. Felids do pose a threat, though Wright (1985) has suggested Aotus are
quite capable of evading them. Larger owls such as the Great-horned owl may pose a threat to the
infants and juveniles (Wright, 1985).

Activity Patterns
Activity patterns are more diverse than was originally thought in Aotus. In Colombia,
Ecuador and most other habitats, the monkeys are entirely nocturnal (Salano, 1995 and
Rosengreen, pers. comm.). In the palm savannas of Paraguay, Argentina and Bolivia, however,
owl monkeys are partly or entirely diurnal (Mann, 1957; Rathbun & Gache, 1977; Wright, 1983).
Aotus is usually more active during the full moon than the new moon (Erkert and Grober, 1986).

Why Be Nocturnal?
Characteristics of Nocturnal Animals
Nocturnal animals, particularly mammals, tend to have small, non-gregarious social
systems. Low light levels require an increase in reliance on auditory and olfactory means by which
to communicate (Charles-Dominique, 1974). Olfaction is probably more effective at night due to
higher olfactory transmissions rates (Wright, 1995). Color vision tends to be lacking. Nocturnal
species tend to be small, as studies in Gabon and Barro Colorado Island in Panama have shown:
both birds and mammals have much lower body weights than do diurnal. No nocturnal primate

exceeds 1 kg (Charles-Dominique, 1974) and no nocturnal Neotropical mammal exceeds 6 kg

(Wright, 1995).
Olfaction is the primary sense in locating plant foods (Charles-Dominique, 1977).
Insectivory tends to be a more important component of the diet in nocturnal animals than in diurnal.
Capturing insect prey requires good hearing and dexterity (Charles-Dominique, 1974).

It is

worthy to note no diurnal strepsirhines save Lemur catta are predatory (Petter, 1962).

Three Hypotheses for the Development of Nocturnality in Aotus

Aotus shares all of the aforementioned characteristics of nocturnal animals. However, the
question of why nocturnality is beneficial to the owl monkeys reproductive success remains
unclear. Nocturnality was not the default character state in Aotus, rather a derived trait
developed from a diurnal ancestor. Lacking a tapetum lucidum, Aotus does not have the overly
specialized eyes for night living. Still, the success of the owl monkey at colonizing much of the
neotropics is undeniable. Three explanations present themselves to the reasons in which this
unique shift in activity pattern evolved: response to interspecific competition for resources,
exploitation of an un- or under-utilized food niche, and avoidance of predators.

Avoidance of Direct Interspecific Competition

The most convincing explanation as to the benefits of nocturnality is due to the avoidance
of interspecific competition. Aotus has little direct contact with diurnal animals who might displace
them from their feeding sites. One of the most important pieces of evidence to support this
hypothesis comes in the differences in feeding tree size between Aotus and Callicebus, a similar
diurnal species. Callicebus, a small monogamous frugivore, exploits smaller-crowned trees which
may not provide enough food or be found quickly enough by larger-bodied and larger-grouped
primates. Larger-crowned trees are dominated by larger primates who will displace Callicebus if
found feeding. There have been reports of Ateles paniscus, Cebus apella and Cebus albifrons
displacing smaller primates from feeding sites.

Saimiri uses large group size to prevent

interspecific displacement by larger primates from becoming effective at keeping them out of larger
fruiting trees. Unlike Saimiri, Callicebus does not have the advantages in numbers (Wright,
1995). Aotus, on the other hand, has little interspecific interaction with larger animals at feeding
sites as most nocturnal animals are both small and solitary (Wright, 1985). During a full moon,
however, Wright (1985) did report displacement by Saimiri and Cebus. Aotus prefers largercrowned trees which can be defended against conspecifics and provide more than adequate
nourishment while allowing for smaller territories. In Paraguay there is a reduced number of
diurnal primates (Wright, 1985) which, by using the hypothesis of interspecific competition
avoidance may explain the diurnal activity patterns of Aotus in this habitat.
Aotus are displaced by Didelphis sp. (opossum), P. flavius, and Bassaricyon sp. but are
never aggressively chased from their feeding site (Wright, 1985).

Little direct or indirect

competition is found with the chiropterans (Estrada et al., 1984), which are not large enough to
pose a threat (Wright, 1985). Lack of intense folivory precludes any intense interactions with
other folivores such as the bamboo rat (Wright, 1985).
By avoiding most sympatric species, Aotus is able to exploit a wider range of habitats
while minimizing dangers of injurious encounters with larger primates and the need to defend large
territories. Being one of the largest nocturnal frugivores (Wright, 1995), the owl monkey is able
to out-compete most sympatric species active at night.

Exploitation of Underutilized Food Niches

Due to high levels of sympatry between many primates, different strategies have evolved to
decrease the niche overlap: species who can exploit underutilized resources with little interspecific
competition will have a selective advantage over those who try to directly compete with sympatric
species (MacArthur, 1958). Aotus uses its nocturnal activity patterns as well as a generalist diet to
minimize the overlap.

A greater reliance on insectivory in Lagothrix than Ateles has been

suggested to be due to microsympatry in fruit diets (Di Fiore, 1997). Aotus eats far more insects

than does Callicebus. Many of the larger insects call and are active at night, making them easier to
locate and confer an advantage to nocturnal insect foraging.
During periods of scarcity, Aotus was able to exploit figs and nectar producing plants,
which Callicebus was not able to due to interspecific displacement by larger-bodied and largergrouped primates. High levels of interspecific competition have also been reported between Ateles
paniscus, Alouatta seniculus and Cebus apella during periods of high fruit productivity
(Guillotin et al., 1994). Aotus rarely ate leaves when other higher-quality foods were available.
However, in the dry forests of Paraguay where seasonally figs and other foods were scarce, Aotus
did become folivorous (Wright, 1995) as well as diurnal (Wright, 1985). Diurnal increases in leaf
proteins and sugars (Ganzhorn & Wright, 1994) and may explain the diurnal activity of Aotus in
Paraguay as an adaptation to maximize the quality of the leaves ingested.

Avoidance of Predators
Predation has historically been a difficult subject to study due to low predator density and
low chances of witnessing a predatory attack.

Predation certainly does have an effect on

populations but the degree to which they can cause major evolutionary responses is, in the opinion
of the author, unlikely to be a major factor in determining the course of evolutionary change.
However, a case can be made for the reinforcement of nocturnality in Aotus being in part
due to the benefits of avoiding predation. The assumption being made is that predation pressures
are less at night in the neotropics than during the day. Wright (1995) has suggested predation by
diurnal raptors is more significant than are nocturnal species. One of the significant problems with
this hypothesis is the fact that while the raptors are diurnal, many other predators are nocturnal
including the snakes and carnivores (Heymann, 1995). It is true that raptors rarely, if ever, prey
on Aotus as the nesting sites are often inaccessible to avian predators. Threats from harpy eagles,
Crested eagles, Ornate hawk-eagles and others (Robinson, 1994) are significant, but the effects on
diurnal monkeys has been as yet unquantified.

Evidence to support the predation hypothesis includes the cryptic coloration of the fur and
the relatively loud vocalizations used in communication which would attract predators if they were
in proximity (Moynihan, 1976). Male parental care may be a defense against predation (Wright,
1984) as males tend to be more vigilant than females (van Schaik and Horstermann, 1994) but is
more likely due to intraspecific defense against infanticide by solitary males and alleviation of the
energetic costs of child-rearing on the mother. Cathemerality in certain populations of Aotus has
been suggested to be due to the lack of diurnal predators (Wright, 1989; van Schaik and Kappeler,
1996) and the existence of large owls (Wright, 1995).

This is further supported by lack of

predictable sleeping sites or regular travel routes in these areas (Wright, 1995). Sleeping sites
often are well-hidden, contain escape routes and offer a high degree of visibility (Aquino and
Encarnacion, 1986).

Conclusions
Reductionist, Apollonian viewpoints as to the reasons behind nocturnality in Aotus are
self-defeating. Rather, a more holistic, multi-causal explanation must be proposed if the system is
to be fully understood. Many features of Aotus are adaptations to a nocturnal lifestyle but are not
necessarily the reasons behind the development of the behavior.

Other explanations may be

important in differing degrees and not mutually exclusive of one another.

Aotus benefits from a nocturnal lifestyle through avoidance of interspecific competition and
predation. Food resources are probably the most limiting factor in Aotus reproductive success.
The social system is unique in that males have a much more equal parental investment. Without a
male, the female is probably completely unable to take care of the infant. Small-bodied primates
require higher quality foods in their diet, being unable to effectively digest enough low-quality
foods to sustain life. The important factor in competition, therefore, is over food resources. The
ability to utilize and defend large concentrations of high-quality foods requires large body size,
large groups or nocturnality. Aotus resides in large-crowned trees and rarely face interspecific
competition, except with the occasional marsupial or one of the few other large mammals in which

little if any aggression occurs (Wright, 1995).

The only significant competition occurs

intraspecifically, as Aotus territories often have a high degree of overlap (Wright, 1978). Very
large fruiting trees often contain multiple groups of owl monkeys displaying intense agonism
towards conspecifics (Wright, 1981).
Predation are probably less important than foraging pressures in determining degrees of
influence as causal mechanisms by which nocturnality evolved in Aotus.

Unfortunately, little

quantitative work has been done to examine the intensity of predation on either the owl monkeys or
their diurnal counterparts. Predation is probably rare in either nocturnal or diurnal primates.
However, Aotus hypothetically lower predation rates may be viewed as an added benefit of being
nocturnal rather than an adaptive response to high diurnal predation leading to a shift in activity
patterns. The evidence to either support or deny the degree by which predation influences the
evolution of any population is scarce and often anecdotal. The author does not deny the potential
influence of predation on primate populations, but further research needs to be done before any
concrete conclusions can be inferred.
Broader-ranging implications of Aotus lifestyle lend support to MacArthurs (1958)
competitive exclusion hypothesis. Primates tend to be broadly sympatric but various life history
patterns show a more specific divergence in strategies (e.g., Di Fiore, 1997). Aotus is an excellent
species to examine this concept due to a high degree of diet-sharing with other primates and its
highly unique activity pattern. Geographical variations on life history patterns can also be
examined. Aotus, ranging across vastly different ecosystems, can yield insights into intraspecific
variation in behaviors. Food choice is another important topic which can be examined using Aotus
as a model. Typically frugivorous and insectivorous, Aotus is able to exploit leaves in scarce
seasons and regions. How Aotus is able to subsist on low-quality foods while being a smallbodied primate suggests some interesting issues of foraging strategies as well as possible circadian
cycles in leaf nutritional content (Ganzhorn and Wright, 1994). The uniqueness of Aotus lifestyle
may prove invaluable in utilizing the comparative method to discuss these broader ideas.

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