Wied's Marmoset (Callithrix kuhli)

MORPHOLOGY: Wied's marmoset has non-opposable thumbs and nails of the digits, which are more claw-like. The crown of this species black along with the ear-tufts (Stevenson and Rylands, 1988). Around the cheeks and the brow there is an area of gray-white coloration (Stevenson and Rylands, 1988). The body pelage is annulated black-gray-orange, with an annulated black-gray tail (Stevenson and Rylands, 1988). Wied's marmoset has a median nasal white stripe (Groves, 2001). The young have juvenile pelage that starts to change into adult pelage around 5 months of age (Stevenson and Rylands, 1988). Infants will start growing ear-tufts at two weeks (Stevenson and Rylands, 1988). The dental formula for this species is 2:1:3:2 on both the lower and upper jaws (Ankel-Simons, 2000). RANGE: Wied's marmoset is endemic to the Atlantic coastal forest of Brazil (Rylands et al., 1993). Wied's marmoset ranges between the Rio Jequitinhonha to the north and the Rio de Contas to the south in the state of Bahia (Mittermeier et al., 1982; Rylands et al., 1988). This species was found to live in evergreen hygrophylic forest, young and old secondary forests, in abandoned rubber plantations, restinga forest, and piacava forest (Stevenson and Rylands, 1988; Rylands et al., 1988; Rowe, 1996). ECOLOGY: Wied's marmoset is a frugivore-insectivore, which also consumes flowers, exudates, nectar, lizards, snails, and frogs (Stevenson and Rylands, 1988; Rylands, 1989; Raboy et al., 2006). Plant galls have also been observed to be eaten by this species (Rylands, 1982, 1989). Grasshoppers were found to be the most preferred insect prey; these include the short-horned (Acrididae) and the long-horned (Tetigoniidae) types (Rylands, 1982). Insects disturbed by army ant, Eciton burchelli, raids are exploited (Rylands, 1982). This species has also been observed to eat stick insects, Phibalosoma phyllinum, (Rylands, 1982). Wied's marmoset also feeds on small lizards and frogs, eating the head first (Rylands, 1982). Snails are eaten by first biting off the apex of the shell and then removing the snail (Rylands, 1982). Small fruit is consumed by removing the fruit with either hands or mouth and eaten while being held in the hand, with the seeds and skin sometimes being discarded (Stevenson and Rylands, 1988). Large fruits are eaten from the branch (Stevenson and Rylands, 1988).

In an area of low second growth forest and scrub forest in an abandoned rubber plantation, Rylands (1982) found that flowers of Symphonia globulifera and the fruit of Dialium guianense were important food sources. Wied's marmoset also fed extensively on fruits and flowers of Melastomataceae (Rylands, 1982). Rylands (1982) also found that exudates were fed upon extensively, especially those of the species Parkia pendula. Frugivory comprised 63% to 70% of feeding and folivory and gummivory comprised 30% to 34% of feeding (Rylands, 1982). Ferrari and Rylands (1994) found that exudates comprised 33.8% of plant parts eaten and fruit, seeds, nectar, and flowers comprised 66.2% of plant parts eaten.

Wied's marmoset feeds on exudates by anchoring the upper incisors in a fixed position on the bark, and uses the lower dentition to scoop out the bark (Coimbra-Filho and Mittermeier, 1978). Exudates are consumed by licking or scooping with the teeth (Coimbra-Filho and Mittermeier, 1978). Exudates include both gum and sap that is eaten by Wied's marmoset (Stevenson and Rylands, 1988). Rylands (1982) observed Wied's marmoset consuming exudates from the following plant species: Parkia pendula, Inga sp., Henriettea succosa, Cardoluvica sp., Sloanea sp., Cupania sp., Trichilia lepdota, and Piptadenia contorta. Parkia pendula was found to be the most important source of exudates (Rylands, 1989). This species will scent mark all of the holes of which they gouge with the circumgenital gland (Stevenson and Rylands, 1988). Some of the holes, however, were not used for feeding, and the marmosets may have gouged first to help absorb the scent mark with the suprapubic gland; these particular scent marks occurred on horizontal branches that were near the boundaries of the territory where territorial disputes were common (Stevenson and Rylands, 1988). Wied's marmoset was found to be more gummivorous than the sympatric tassel-ear marmoset, Callithrix intermedia (Stevenson and Rylands, 1988).

Rylands (1989) found that when this species forages, the groups would travel slowly, investigating the area thoroughly. The locomotion tends to be stealthy as compared to other types of locomotion, and individuals will be alert and concentrate on their surroundings (Rylands, 1989). This species has been found to capture prey by pouncing and trapping; also it will look under leaves and clumps of debris and investigate holes in branches (Rylands, 1989). Rylands (1989) found that this species spent 78% of the time below 15 meters in the forest (Rylands, 1989). Foraging may predominantly occur in the lower levels of the forest because of the frequent utilization of army ant swarms (Rylands, 1989).

The mean group size for this species is 6.6 individuals per group (Rylands, 1982). Rylands (1982) found a population density of 50 to 68 individuals per square kilometer at Una, Brazil. The day range length ranges between 942-1080 meters per day (Rylands, 1982). The majority of the time (62.8%), this species is found between 6-15 meters high in the forest (Ferrari and Rylands, 1994). Ferrari and Rylands (1994) found that the daily activity budget for Wied's marmoset is 38.1% locomotion, 27.6% foraging for and feeding on animal material, 20.4% feeding on plant material, and 13.9% rest/social activities. This species sleeps in dense vegetation and liana tangles at heights between 5 and 15 meters, and they were found to use 14 different sites in a 10-hectare home range in a three-month period (Rylands, 1982, 1989). Large epiphytic Bromeliaceae and Araceae are also used for sleeping sites (Rylands, 1982). Groups have home ranges that overlap with other conspecific groups (Rylands, 1982, 1989). Raboy et al. (2006) found that secondary forest made up 12.2% of a group's home range, but 54.8% of their core area.

Potential predators of Wied's marmoset include: the tayra (Eira barbara), forest hawks (Micrastur spp.), and coatimundis (Nasua) (Stevenson and Rylands, 1988). Carroll and Schaffner (1998) found that this species has a preference for portable foods, which could be the result of the need for anti-predator surveillance, portable foods allowing the individual to travel easier.

LOCOMOTION:
Wied's Marmoset moves through the forest quadrupedally, but is capable of leaping (Fleagle, 1988). When leaping, individuals will land on both horizontal and vertical branches (Stevenson and Rylands, 1988). This species can leap up to 3 meters (Stevenson and Rylands, 1988). When ascending large vertical branches, Wied's marmoset will use a squirrel-like locomotion, first using the two forelimbs then use the two hindlimbs (Stevenson and Rylands, 1988). Vertical surfaces are descended head first (Stevenson and Rylands, 1988). The tail is held up and swings from side to side when carefully moving along thin vines and branches (Stevenson and Rylands, 1988). The tail is held horizontally, sometimes slightly arched, during rapid quadrupedal locomotion (Stevenson and Rylands, 1988). This species will hang from vertical supports with just the hindlimbs (Stevenson and Rylands, 1988).

SOCIAL BEHAVIOR: The basic family group is composed of the adult pair, their offspring, and sometimes other individuals (Alonso et al., 2000). Within the group, there are two subgroups, the adult pair and the juveniles (Alonso et al., 2000). New groups may form by splitting and dividing the home range (Rylands, 1982). This species has a polyandrous mating system. Adults spend little time in contact with juveniles (Alonso et al., 2000). Alonso et al. (2000) found that the strongest social bond is between juvenile group members, with the second most strongest between the adult pair. The hierarchy within the groups are maintained by agonistic vocalizations directed at a challenger (Alonso et al., 2000). Schaffner and Angel (1996) found that there is a clear hierarchy between related adult males in a group in terms of access to food, higher rates of sexual activity, and exhibiting higher rates of territorial behavior. One male will maintain proximity and have copulatory bouts with breeding females more than other group males (Schaffner and French, 1994). During female maximum fertility, one male will monopolize access to the female (Schaffner and French, 1994). Although recently, Schaffner and French (2004) have found that all group males will engage in sexual activity. This may be the result of sperm competition, weak social bonds between males and females, and the need for multi-individual infant care (Schaffner and French, 2004). In a captive experiment, French et al. (1993) found that males direct higher rates of agonistic threats towards unfamiliar intruders as compared to familiar intruders. Also when exposed to unfamiliar intruders, females will engage in territorial behavior while the males perform mate-guarding behavioral patterns (Schaffner and French, 1995).

All group members care for offspring, carrying and warming them, sharing food, and protecting them from danger (Santos et al., 1997; Angel and French, 1996). Infant carrying by the mother peaks at 1-2 weeks after birth, when androgen levels rise, facilitating the reduction in care (Nunes et al., 2001; Santos et al., 1992; French et al., 2004). Carrying by the father was found to peak at about 3-4 weeks after parturition, declining afterwards, and testosterone and oestradiol levels in fathers decreased when carrying increased (Nunes et al., 2000, 2001). This decrease in hormone levels occurs regardless of whether the infants survive or not (Nunes et al., 2000). The decline in testosterone was found to be greater after the birth of a second litter as compared to after the first litter (Nunes et al., 2001; Puffer and French, 2004). When young are weaned, testosterone and oestradiol levels increase significantly in males (Nunes et al., 2000). Fite et al. (2004) found that in females an increase in testosterone results in a decrease in maternal effort. The amount of carrying by the father is consistent from litter to litter (Nunes et al., 2001). Santos et al. (1997) found that carrying by the father was more in small groups as compared to large groups during early infant development. The opposite is true with mothers, in large groups carrying percentage is higher than in small groups (Santos et al., 1997). Carrying by siblings peaked from 1-4 weeks after birth, but was found to be overall low in occurrence (Nunes et al., 2001). Infant carrying may be influenced by a small home range size, short ranging distance, and slow travel speed (Santos et al., 1997). Infant mortality of this species and other callitrichids is higher in groups with one or no alloparents to assist the parents with carrying (Santos et al., 1997). Infant carrying is also dependent on whether the infant is chimeric, where mothers have a tendency to carry chimeric infants less while fathers tend to carry chimeric infants more than non-chimeric infants (Ross et al., 2006) Adults with young were found to spend less time sleeping throughout the night than adults without young (Fite and French, 1999; Fite et al., 2002).

Social play was found to occur most frequently amongst young marmosets (Alonso et al., 2000). Solitary play is confined to young individuals (Alonso et al., 2000).

During intergroup encounters, individuals will chase each other, ending with two individuals facing each other while showing tail raised present, emitting twitter, piloerection, and tuft flick (Rylands, 1982). Encounters rarely involve fights (Rylands, 1982). Males will exhibit more aggression than females, especially when presented with unfamiliar intruders, and males will perform more agonistic display towards unfamiliar intruders than familiar intruders (French et al., 1995). Males have been found to approach familiar intruders more than females (French et al., 1995). Female-female aggression may vary between groups, with some females showing more aggression (French et al., 1995).

Wied's marmoset may form mixed-species groups with Leontopithecus chrysomelas, and are able to maintain the groups because they do not compete for the same food resources and forages in different niches (Rylands, 1989; Raboy and Dietz, 2000). Leontopithecus chrysomelas forages in the upper story of the forest and Wied's marmoset forages in the middle to lower stories (Rylands, 1989). Mixed-species groups are maintained with loud calls (whirrs) (Raboy and Dietz, 2000). This species has the smallest mean group size (6.6) for marmosets (Rylands, 1982, 1989).

VOCAL COMMUNICATION:
whirr: This is a repeated element of varying length at about 8 kilohertz (Stevenson and Rylands, 1988). This call consists of a cyclically fluctuating frequency (Stevenson and Rylands, 1988). This call is very common (Stevenson and Rylands, 1988). This is used as a contact call, intragroup (Stevenson and Rylands, 1988). All individuals emit this call (Stevenson and Rylands, 1988). The motivation is to maintain group contact (Stevenson and Rylands, 1988; Schaffner et al., 1995). The response is to reciprocate with the same call (Stevenson and Rylands, 1988).

long call: This call has a loud an repetitive morphology (Jorgensen and French, 1998; Jorgensen, 1996). This is also called a phee call, where there are three to seven phee vocalizations in a row (Schaffner et al., 1995). This is emitted by both adult males and females (Schaffner et al., 1995). This call is distinct amongst individuals and is sexually dimorphic (Jorgensen and French, 1998; Smith et al., 2007). This call shows a lack of stability over time (Jorgensen and French, 1998). Rukstalis et al. (2002) suggests that this call is an example of maintaining vocal plasticity beyond the infant stages.

trill: This serves as a contact call (French et al., 1995). This call is emitted increasingly when a pair is presented with a familiar or unfamiliar male intruder (French et al., 1995). This call is also emitted increasingly with the presentation of an unfamiliar female (French et al., 1995).

OLFACTORY COMMUNICATION:
suprapubic marking: This is when an individual presses the suprapubic pad against a substrate and deposits secretions by pulling itself along or by pushing itself with its feet (Epple et al., 1993; Schaffner et al., 1995). This behavior pattern is frequent in Wied's marmoset and can occur more frequently by an individual where the home ranges of two groups overlap, thus suggesting a use for demarcation of territory (Rylands, 1990).

anogenital marking: Individuals will perform this at higher rates during feeding times (Forman and French, 1992).

VISUAL COMMUNICATION:
bouncing gait: This is an individual runs having an exaggerated bouncing movement (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This behavior is seen during situations of play and in locomotion by infants (Stevenson and Rylands, 1988). Young individuals perform this display (Stevenson and Rylands, 1988). This functions to initiate play and is motivated by the need to play or by exploration (Stevenson and Rylands, 1988). The response to bouncing gait is that the individual may be played with by the receiver (Stevenson and Rylands, 1988).

gallop: This is when an individual runs quickly; the tail may be extended during this behavior (Stevenson and Rylands, 1988). This behavior is described as being very common in occurrence (Stevenson and Rylands, 1988). This is seen during play and running away (Stevenson and Rylands, 1988). This is performed by all individuals (Stevenson and Rylands, 1988). This serves to initiate or respond to play, or this serves to flee from an aggressor or predator (Stevenson and Rylands, 1988). This has both playful and aggressive or fearful motivations (Stevenson and Rylands, 1988).

stalk: This is when one individual will visually fixate on another whilst hiding, and makes jerky movements towards the stimulus (Stevenson and Rylands, 1988). This is a common behavior (Stevenson and Rylands, 1988). This is seen in situations of play and the ontogeny of play (Stevenson and Rylands, 1988). This is performed by all individuals (Stevenson and Rylands, 1988). This serves to communicate the initiation of play and has playful motivations (Stevenson and Rylands, 1988). The response is that the sender may be played with by the receiver (Stevenson and Rylands, 1988).

slide: This is when an individual moves on the side, and is propelled by the arms and legs (Stevenson and Rylands, 1988). This behavior is seen during social and solitary play (Stevenson and Rylands, 1988). This is said to be performed by offspring (Stevenson and Rylands, 1988). This has playful motivations and serves as a response to play initiation (Stevenson and Rylands, 1988).

roll: This is when an individual rolls onto the back or side, and includes a somersault (Stevenson and Rylands, 1988). This is a common behavior (Stevenson and Rylands, 1988). This is seen in situations of social and solitary play (Stevenson and Rylands, 1988). This is said to be performed by offspring (Stevenson and Rylands, 1988). This has playful motivations and serves as a response to play initiation (Stevenson and Rylands, 1988).

leg stand: This is when an individual stands on its hind-legs with the hands outstretched (Stevenson and Rylands, 1988). This behavior is rare (Stevenson and Rylands, 1988). This is seen when the individual is observing an object or a conspecific (Stevenson and Rylands, 1988). This is performed by all individuals (Stevenson and Rylands, 1988). This display functions as being a non-response or response to aggression and is motivated by interest and fear (Stevenson and Rylands, 1988). This also may be performed to gain better visibility (Stevenson and Rylands, 1988).

withdrawal gesture: This is when an individual withdraws the body and arms remain fully or partially extended (Stevenson and Rylands, 1988). This behavior is rare (Stevenson and Rylands, 1988). This is seen during situations of conflict (Stevenson and Rylands, 1988). This is performed by all individuals except the adult pair (Stevenson and Rylands, 1988). This display functions to communicate submission and is motivated by fear (Stevenson and Rylands, 1988). The response is that the receiver may withdraw or attack (Stevenson and Rylands, 1988).

cringe: This is described as a more extreme form of withdrawal gesture, but having the hind-legs bent (Stevenson and Rylands, 1988). This behavior is rare (Stevenson and Rylands, 1988). This is seen during situations of conflict (Stevenson and Rylands, 1988). This is performed by all individuals except the adult pair (Stevenson and Rylands, 1988). This display functions to communicate extreme submission and is motivated by extreme fear (Stevenson and Rylands, 1988). The response is that the receiver may withdraw or attack (Stevenson and Rylands, 1988).

slow tongue in/out: This is when the tongue is moved in and out rhythmically (Stevenson and Rylands, 1988). This behavior is rare (Stevenson and Rylands, 1988). This is seen during grooming (Stevenson and Rylands, 1988). This is performed by all individuals (Stevenson and Rylands, 1988). This behavior is seen sometimes during grooming, but not always (Stevenson and Rylands, 1988).

head cock stare: This is when an individual moves the head from side to side while observing the stimulus (Stevenson and Rylands, 1988). This is seen when an individual is observing an object or conspecific (Stevenson and Rylands, 1988). This is performed by all individuals (Stevenson and Rylands, 1988). This functions to aid vision and is motivated by interest (Stevenson and Rylands, 1988).

open mouth: This is when an individual has the mouth open with the teeth sometimes showing (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during play bouts, often combined with other play patterns (Stevenson and Rylands, 1988). All individuals perform this display, though it tends to be used more by adult males (Stevenson and Rylands, 1988). This serves to metacommunicate play and is motivate by playful intentions (Stevenson and Rylands, 1988). The response is that the receiver may play or not (Stevenson and Rylands, 1988).

rhythmic open mouth: This is when the mouth is opened and closed rhythmically, directed towards opposite-sexed individuals (Shepherd et al., 1992). Either sex will initiate this behavior, and is followed by the male mounting and copulating with the female (Shepherd et al., 1992). Santos and Otta (1996) found that this display is used to coordinate reproductive behavior between partners and for the male to inform the infant of danger so the infant climbs on the back.

slit stare: This is when an individual stares having the eyelids half closed, the eyes in a slit (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during agonistic situations and less commonly during play (Stevenson and Rylands, 1988). All individuals perform this display (Stevenson and Rylands, 1988). This is a response to being approached by an aggressive conspecific or when approaching a strange object (Stevenson and Rylands, 1988). This display has defensive motivations (Stevenson and Rylands, 1988).

frown: This is when an individual lowers the eye brows (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during agonistic situations (Stevenson and Rylands, 1988). All individuals perform this behavior (Stevenson and Rylands, 1988). This serves to cause an approaching conspecific to retreat and is motivated by mild aggression (Stevenson and Rylands, 1988). The response is that the receiver may retreat (Stevenson and Rylands, 1988).

partial open mouth: This is when the middle portion of the mouth is open and the teeth visible (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen when approaching a strange object or during an agonistic situation (Stevenson and Rylands, 1988). All individuals perform this display (Stevenson and Rylands, 1988). This is a response to being approached by an aggressive conspecific or when approaching a strange object (Stevenson and Rylands, 1988). This is motivated by fear (Stevenson and Rylands, 1988).

bared-teeth gecker: This is where the lips are retracted, middle portion of the mouth is open, and the teeth are visible (Stevenson and Rylands, 1988). This is accompanied by a gecker vocalization (scream) (Stevenson and Rylands, 1988). This behavior is very common (Stevenson and Rylands, 1988). This is seen during agonistic situations (Stevenson and Rylands, 1988). This is performed by any subordinate individual (Stevenson and Rylands, 1988). This display is performed by a subordinate during a fearful situation and has submissive motivations (Stevenson and Rylands, 1988).

bared-teeth scream: The lips are retracted more and the mouth open wider as compared to bared-teeth gecker (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during agonistic situations (Stevenson and Rylands, 1988). All individuals perform this display (Stevenson and Rylands, 1988). This display is performed by an individual who is being attacked by another, and it has fear and submissive motivations (Stevenson and Rylands, 1988).

tufts/ear flick: This is when an individual will flick the tufts/ears back and forth (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This behavior occurs in agonistic situations, but not exclusively (Stevenson and Rylands, 1988). All individuals perform this behavior (Stevenson and Rylands, 1988). This is a submissive response to agonistic situations and is motivated by mild aggression (Stevenson and Rylands, 1988).

tufts/ears flatten: This is where the tufts/ears are flatten against the head (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during agonistic situations and during play (Stevenson and Rylands, 1988). All individuals perform this display (Stevenson and Rylands, 1988). This is a submissive response to agonistic situations, and in these situations it is motivated by fear (Stevenson and Rylands, 1988).

tufts/ears forward: This is where the tufts/ears are held forward with the face possibly reddening (Stevenson and Rylands, 1988). This behavior is rare (Stevenson and Rylands, 1988). This behavior is seen during aggressive situations and may precede an attack (Stevenson and Rylands, 1988). All individual perform this display (Stevenson and Rylands, 1988). This serves to communicate aggression and has aggressive motivations (Stevenson and Rylands, 1988). The response to this is to either retreat or attack on the part of the receiver (Stevenson and Rylands, 1988).

arch bristle locomotion: This is when an individual will strut with the back arched and the pelage erect (Stevenson and Rylands, 1988). The pelage of the tail is more erect at the base (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during intergroup display and less frequently during intragroup displays (Stevenson and Rylands, 1988). All individuals over 10 months perform this display (Stevenson and Rylands, 1988). This is used in ritualized display with other group members and has agonistic motivations (Stevenson and Rylands, 1988). The response to this is that another group will reciprocate (Stevenson and Rylands, 1988).

pilo-tail: This is when part or all of the tail pelage is erect (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen in situations of alarm (Stevenson and Rylands, 1988). All individuals perform this behavior (Stevenson and Rylands, 1988). This behavior has fearful motivations (Stevenson and Rylands, 1988).

tail raised present: This is when the tail is semi-piloerected, raised, and may be coiled (Stevenson and Rylands, 1988). The genitals are exposed (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during aggressive situations directed at another individual (Stevenson and Rylands, 1988). All individuals over 8-10 months perform this display (Stevenson and Rylands, 1988). This has a function during intergroup ritualized displays and during intragroup aggression (Stevenson and Rylands, 1988). This display has agonistic motivations (Stevenson and Rylands, 1988). When used during intragroup aggression, the response is usually a submissive response (Stevenson and Rylands, 1988).

TACTILE COMMUNICATION:
face/body nuzzle: This is when one individual nuzzles the face or body of another (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen in situations of low activity (Stevenson and Rylands, 1988). All individuals perform this behavior, although it is common between older and younger individuals (Stevenson and Rylands, 1988). This is used to communicate amicable behavior (Stevenson and Rylands, 1988). The response is a vocalization or reciprocal amicable behavior (Stevenson and Rylands, 1988).

copulate: This is when a male mounts the back of the female, having the back legs on a substrate, and showing several bouts of thrusting (Stevenson and Rylands, 1988). The male's tail may be coiled (Stevenson and Rylands, 1988). This behavior is rare (Stevenson and Rylands, 1988). This is seen during mating, but is different than actual mating (Stevenson and Rylands, 1988). This is done most commonly by the adult pair (Stevenson and Rylands, 1988). This is used to bring about copulation and has sexual motivations (Stevenson and Rylands, 1988). The female response is normally amicable (Stevenson and Rylands, 1988).

play bite: This is when an individual will give an inhibited bite to another's body (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during situations of playful interactions (Stevenson and Rylands, 1988). All individuals perform this behavior (Stevenson and Rylands, 1988). This used to initiate play and has playful motivations (Stevenson and Rylands, 1988). The response is to play or not to play (Stevenson and Rylands, 1988).

grooming: This is when one individual parts the hair of another with hands to remove particles with the teeth (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen in situations of low activity (Stevenson and Rylands, 1988). All individuals perform this behavior (Stevenson and Rylands, 1988; Alonso et al., 2000). Alonso et al. (2000) found that in a group the adult male and a young male were the most active groomers. This behavior may elicit more grooming (Stevenson and Rylands, 1988).

pounce: This is when one individual pounces rapidly on another (Stevenson and Rylands, 1988). This behavior is common (Stevenson and Rylands, 1988). This is seen during play and is performed by all individuals (Stevenson and Rylands, 1988). This is used to initiate a play bout and has playful motivations (Stevenson and Rylands, 1988).

REPRODUCTION:
Wied's marmoset gives birth to twins, which also occurs on average in captivity (Fleagle, 1988; Ross et al., 2007). Twinning may have been possible in callitrichines because of the use of exudates as a food source, but a decrease in size and the use of male helpers in caring for young may also have contributed (Ah-King and Tullberg, 2000). Marmoset twins are dizygotic, and could be fathered by more than one male (one twin by one male and the second by another male) (Wislocki, 1939; Stevenson and Rylands, 1988). Males reach reproductive maturity at age 12 months (French and Schaffner, 1995), and females at 12-15 months of age (Smith et al., 1997). Gestation lasts 4.5 months and the interbirth interval is about 5 months (French et al., 1996; Ross et al., 2007). In a group there is one breeding adult female, but subadult females have been found to ovulate while still living with family groups (Smith et al., 1997). Puffer et al. (2004) found that subadult females do have their breeding hormonally suppressed following the breeding female's peripartum period or the return to fertility. Ovulation of females was found to be different when they lived in their natal group versus being kept alone, where in the natal group the ovarian cycles had a short luteal phase and a lower peak concentration of progesterone metabolite excretion (Smith et al., 1997, 1995). Fite and French (2000) found that when there is increased prepartum estradiol levels, it increases the chance of infant mortality.

During courtship males will direct arch-bristle towards coy females (Stevenson and Rylands, 1988). Rylands (1982) found that copulations occurred for one group in October and for another group in November. After first pairing, sexual behavior between the male and female have been found to peak initially, then decrease over time, while allogrooming and grooming solicitation increases over the longevity of the pair-bond (Schaffner et al., 1993, 1995). Initially when the pair-bond is formed, males will play a larger role in maintaining social proximity, but over time both males and females will play an equal role (Schaffner et al., 1993, 1995). Individuals will show elevated sexual behavior when separated from pair-mates for a temporary period of time (Shepherd and French, 1999). Schaffner et al. (1995) found that when a sexual interaction was preceded by rhythmic open mouth, ejaculation was more likely to occur when a sexual interaction was not preceded by rhythmic open mouth.

REFERENCES:
Ah-King, M. and Tullberg, B.S. 2000. Phylogenetic analysis of twinning in callitrichinae. American Journal of Primatology. Vol. 51, 135-146.

Alonso, C., Porfirio, S., and Langguth, A. 2000. Social interactions in a Callithrix kuhlii family (Primates: Callitrichidae) in captivity. A Primatologia no Brasil. Vol. 7, 23-33.

Angel, L.J. and French, J.A. 1996. Infant carrying patterns vary with group demography in black tufted-ear marmosets (Callithrix kuhli). (Abstract) Abstracts of the XVI Congress of the International Primatological Society and the XIX Congress of the American Association of Primatologists. Madison, WI, 691.

Ankel-Simons, F. 2000. Primate Anatomy. Academic Press: San Diego.

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Nunes, S., Fite, J.E., Patera, K.J., and French, J.A. 2001. Interactions among paternal behavior, steroid hormones, and parental experience in male marmosets (Callithrix kuhlii). Hormones and Behavior. Vol. 39, 70-82.

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