range of other sounds, including barks, grunts, woofs, cackles and screeches. It uses clucking sounds to maintain auditory contact with other members of its group. The mantled howler also uses non-vocal communication, such as "urine rubbing" when in a distressful social situation. This consists of rubbing the hands, feet, tail and/or chest with urine. It marks its scent by rubbing its throat on branches. Lip smacking and tongue movements are signals used by females to indicate an invitation to mate. Genital displays are used to indicate emotional states, and group members shake branches, which is apparently a playful activity. The mantled howler is usually indifferent to the presence of humans. However, when it is disturbed by people, it often express its irritation by urinating or defecating on them. It can accurately hit its observers despite being high in the trees. Tool use The mantled howler has not been observed using tools, and prior to 1997 no howler monkey was known to use tools. However, in 1997 a Venezuelan red howler (Alouatta seniculus) was reported to use a stick as a club to hit a Linnaeus's two-toed sloth, (Choloepus didactylus), that was resting in its tree. This suggests that other howlers, such as the mantled howler, may also use tools in ways that have not yet been observed. Reproduction The mantled howler uses a polygamous mating system in which one male mates with multiple females. Usually, the alpha male monopolises the breeding opportunities, but if the alpha male is distracted, a lower-ranking male can get an opportunity to mate. And in some groups, lower-ranking males do get regular mating opportunities and do sire offspring. Alpha males generally maintain their status for about to 3 years, during which time they may father 18 or so infants. Females become sexually mature at 36 months, males at 42 months. Females reaching sexual maturity are typically 42 months old by their first birth. They undergo a regular estrus cycle, with an average duration of 16.3 days, and display sexual skin changes, particularly swelling and color change (from white to light pink) of the labia minora. The copulatory sequence begins when a receptive female approaches a male and engages in rhythmic tongue flicking. The male responds with the same tongue movements before the female turns while elevating her rump, which allows for mating to begin. Females apparently also use chemical signals, since males smell the females'