ABSTRACT Field observation and of sound recording of Formosan macaques (Macaca cyclopis) at Mt. Longevity were conducted from Oct. 1999 to Oct. 2000. Spectrographic analyses revealed at least 25 basic patterns with 32 different vocal types from a total of 375 hrs recordings (55 hrs for scan sampling and 320 hrs for focal animal sampling). The three most frequent calls in overall vocal repertoire were coo calls (18.59%), hack (17.78%) and contact rattle (11.85%). The other calls included squeak (11.34%), noise and undulated scream (6.61%), sneeze (4.54%), greeting (3.77%), squeal (3.41%), vibrato growl (3.02%), growl (2.86%), cluck (2.54%), female copulation call (2.45%), squawk (2.14%), tonal scream (2.03%), alarm call (1.74%), threat rattle (1.66%), chuckle (0.92%), rise (0.61%), weeping (0.56%), male copulation call (0.32%), babble (0.30%), whine (0.29%), roar (0.27%), bark (0.24%) and mounting grunt (0.15%). Formosan macaques employ a complex vocal system composed of discrete signals as well as graded signals which vocal patterns connected by intermediate gradations. Discrete signal included coo call, greeting, contact rattle, mounting grunt, two copulation calls, babble, whine, weeping. Whereas the graded signals included the aggressive signals (threat rattle, growl, bark, roar, vibrato growl and alarm call), chuckle, the submissive signals (noise and undulated scream, squeal, tonal scream and squeak) and the distress signals (squawk, hack, cluck and sneeze). According to the context of emission, some of vocalizations could be divided into: (1) Affiliative contact calls: The calls, as the coo calls, greeting and contact rattles, may be summarized as affinitive contact calls for reduction and/or maintenance of close proximity between group members. (2) Aggressive calls: The growls, threat rattles and vibrato growls were uttered by dominant animals menacing sub-dominate group mates, members of other groups or other species. (3) Submissive calls: these submissive calls (including the various form of scream, squeal and squeak) with obviously structure differences. Male and female copulation calls and alarm call of M. cyclopis were clearly related to particular interactions or specific external stimulus. The acoustic structures might be influenced by specific social factors, motivation or the arousal state of the callers. The acoustic structure of submissive calls was associated with the aggression with or without physical contact. Noise and undulated scream were usually used in aggressive interactions with physical contact (68%), whereas squeal, tonal scream and two types of squeak were used without physical contact (75%). Age specific difference of vocal behavior seemed to more pronounced than asymmetries between the sexes. Infants have the highest vocal rate (1142.2 vocalizations/10 h) and relative frequency of vocalization (44.58 %) than other age/sex classes and they decreased from infants to adults. With increasing age, visual signals could not only complement vocal signals but also replace the vocal cues. The relative frequency of four vocal patterns (coo call, greeting, squeak and hack) decreased from infants to adults, whereas that of threat rattle and growl increased from infant to adults. On the other hand, there were five vocal patterns (including cluck, sneeze, whine, weeping and babble) only used by juveniles and infants. In addition, seven vocal patterns included squeak (64 %), hack (63.6 %), squawk (76%), cluck (85.7 %), sneeze (90 %), weeping (88 %) and babble (100%) mainly were used by infants. Among them, three vocal patterns (squeak, hack and squawk) were produced mostly by infant II. Only infants used babble. Roar and bark were only present in adults and sub-adults. Obvious sex differences of relative frequencies of vocal patterns occurred in adults, the vocal activity of females consistently higher compared to that of males. In six patterns (vibrato growl, chuckle contact rattle, squeal, squeak and hack), females uttered more often than male peers. Mounting grunts and male copulation calls were exclusively used by adult males whereas female copulation calls were only produced by adult/sub-adult females. Vocalizations were produced/ceased by certain sex/age class that might associate with social organization, morphological feature and circumstances experience. The vocal repertoires of Formosan macaques revealed the high similarities in the species within genus Macaca, especially to M. fuscata, M. mulatta and M. radiata. It may be due to phylogeny, habitats and social organizations.

This study investigated the mother-infant relationships of Formosan macaques (Macaca cyclopis) at the Mt. Longevity during the first 24 weeks of infants¡¦ age. The field observation took place from January to November 2002 and from April to December 2003. The total observation time recorded was 450 hours. The death rate of infant males (23.7%) was higher than that of infant females (2.8%). The death rate of infants born at the later period (41.7%) was higher than those of infant born at the earlier and the peak periods (7.7%, 8.2%). The death rate of infants from primiparous females (30.8%, 4/13) was slightly higher than that of infants from multiparous females (9.8%, 6/61, p>0.05). During the observation period, I followed 43 mother-infant dyads, but 5 infants died or disappeared, and only 38 pairs left. Mother¡Vinfant relationships in Formosan macaques were influenced by infant age and sex, matriline size and the number of immature sister of the infant. The percentages of time that mother-infant contact, sucking, mother carrying ,cradle infant, and the percentages of number that contact made by mother and mother restrain infant broken contact were decrease as infants grow older. On the other hand, the percentage of time that mother-infant distance > 1 meter and mother grooming increased as infants older. But mother reject infant contact was not affected by infant¡¦s age. Adult females spent more time carrying female than male infants when infants were one week old. Developments in jumping and eating were seen earlier in male than female infants. The percentages of time in ventro-ventral contact in mother-infant dyads decreased as the number of infants¡¦ immature sisters increased within infants¡¦ first month of age. When a mother wounded, she spent less time in contacts with her infant; however, when the infant wounded, mother¡Vinfant dyads spent more time in contacts. The data provide a better fit to the Reciprocity hypothesis because the percentage of the female (87.3%, 234/268) to take care of infants was higher than male (12.7%). The percentage of the adult female (allomother) to take care of female infants (59.0%, 79/134 ) is higher than taking care of male infants (41.0%, p<0.005 ). The percentage of the adult female that takes care of non-blood related infants (81.6%, 71/87) is considerably higher than the percentage of taking care of blood-related infants (18.4%, p<0.0001). The percentage of adult female that grabs infants roughly (87.4%, 83/95) is higher than juvenile female (7.4%, 7/95 ).

Abstract The present study investigated the activity patterns of Formosan macaques at Mt. Longevity and an emphasis was given to feeding and foraging behavior. The diet composition of Formosan macaque showed significant changes among different age and sex classes of macaques as well as among different seasons. The field research was carried out from August 2003 to July 2004 for a total of 311 hrs covering 77 days. Feeding behavior dominated the activity patterns of the macaques at Mt. Longevity (28.11%), followed by other behaviors such as affiliate (24.71%), resting (17.10%) and moving (16.04%). In contrast, foraging (8.22%) and agonistic behaviors (5.50%) were the least among the activity patterns recorded during this study. Interestingly, adult males spent more time in resting (30.60%) while adult females spent more time in feeding (29.84%), which indicated that the activity patterns were influenced by sex and age groups. Plant food items accounted for 94.87% of their diet while the seminal fluid and breast milk accounted for 5.08%. The plant food items consumed by the monkeys came from 46 species in 31 families and the fruits alone amounted 42.18%. Other food items included leaf (26.20%), stem (11.84%), flower (10.33%), bud (9.14%) and root (0.03%). The relative frequency of fruit consumption by the macaques was higher than 40 % between May and September (40.53% - 63.79%) and December to January (54.66% - 55.28%). The Formosan macaque diet composition changed according to the sex and age class groups. The Simpson index, Shannon-Wiener index and Levine¡¦s niche breadth index changed according to sex, age and months respectively. The plant food diversity indexes were highest for the juvenile males and lowest for the adult females. The Levine¡¦s niche breadth index was broader for the juvenile females but narrowest for the adult females. Adult females consumed 35 species of plants which is higher than the number of species that the adult males and juveniles consumed. However, the plant food diversity index and the niche breadth index were low for adult females, which indicate that they choose certain types of food. On the other hand, all these three indexes were highest in May and changed from month to month. The diet overlap of the Formosan macaque (Renkonen percentage) was highest between adult males and adult females (85.2%) than between other sex/age groups. Among the five categories of age and sex groups, the diet overlap was 66.68% and 22 species of plants was consumed by all age and sex groups of Formosan macaques.

Abstract This study analyzed the alarm calls of Formosan macaques and their functions from Mt. Longevity. Under the natural condition, 137 five-minute scan sampling and 129 completely 20-minute all occurrence sampling were collected to record behaviors and the alarm calls. In addition, 43 stimulus (predator) tests and 87 playback experiments were successfully conducted from September 2002 to March 2003. Incidents that triggered alarm calls included: the confrontation with dogs, threats from travelers with a cane, slingshots, or stones, encounters with other troops of macaques, passing-by motors or mobile cars, airplanes flying above, and some unrecognizable factors. Under the natural condition, most of the alarm calls produced by Formosan macaques were responds toward dogs (63.46%), with the average frequency of 0.78 times/10hrs. In the predator tests, types of stimulus (dog, human with slingshot, m-snake1 and m-snake2), sex/age classes (adult male, adult female, juvenile, and infant), and the position of macaques (0m, <1m, 1m, and ¡Ù2m) had great influence on the behavioral response of macaques (p<0.01). The response scores were from 3 to 0 (move away more than 5 times body length or climbed to tree, move away up to 5 times body length, visual orientation towards the predator and no apparent response). The average response score of macaques to dogs was the highest one, far above threats from a person with a slingshot, m-snake1 and m-snake 2 (p<0.05). When Formosan macaques confront dogs, the vocal frequency (time/individual) to the alarm call had great influence on sex/age classes of macaques (p<0.01), and the average alarm call frequency was highest from adult males. When human with slingshots and m-snake were predators, majority of the adult males and females adopted run-away (69.91¢M), while very few climbed up trees (8.02¢M) to prevent from any harm caused by predators. However, they ran away (31.52%) or climbed up trees (48.47%) in response to dogs in different proportions. The average response score of macaques, from high to low, was from infant, juvenile, adult female and adult male, and the differences were significant (p<0.05). When Formosan macaques encounter these four types of predators, their alarm calls were quite similar in the spectrographs. The six basic vocal characteristics (maximal, median and minimal fundamental frequency, lowest and peak frequency and duration), analyzed by canonical discriminate analysis, indicated that alarm calls of Formosan macaques confront dogs and human with slingshots could be distinguished from snake models. The four types of predators had significant effects at the median and maximal fundamental frequency of the basic vocal characteristics from adult females and juveniles (p<0.01). But there was no difference in the nine basic vocal characteristics of alarm calls from adult males and infants toward four types of predators. The alarm calls of adult females and juveniles toward dogs in the median and maximal fundamental frequency both were significantly lower than those from m-snake1. In the playback experiments, sex/age classes, types of the alarm calls toward stimulus (dogs, travelers with slingshots, m-snake1 and m-snake 2) and the position of macaques had significant effects on their behavioral responses (p<0.01). The average response score of macaques in playbacks, from high to low, was from infant, juvenile, adult female, and the adult male, and the differences were significant (p<0.05). When the alarm call caused by dogs played back, the average response score of macaques was higher than the alarm calls stimulated by m-snake1 and m-snake 2 (P<0.05). In the playback experiments, when macaques at a higher place (¡Ù2m), they often visually orientated towards the predator or no apparent response whit a lowest response score. The average response scores of the adult males and females toward four types of alarm calls (playback) were higher than the control ones (p<0.05).

The purpose of this study was to analyze and compare the agonistic behaviors and reconciliation in captive and free-ranging Formosan macaques (Macaca cyclopis). The dominance style of Formosan macaques was compared with long-tailed, rhesus and Japanese macaques. I have used scan, focal sampling and ad libitum on aggressions of adult macaques. I have recorded post-conflict (PC) focal samplings on victims and compared those with matched control (MC) focal samplings. Agonistic behaviors had significantly higher frequency in captive than in free-ranging Formosan macaques. The frequencies of hostile and submission were significantly higher in captive than in free-ranging Formosan macaques. The captive adult females of higher rank had higher frequency of threat and hostile, and lower frequency of submission. Threat was the most frequent aggression (52-72%) expressed by both the captive and free-ranging adult monkeys. The victims in captive and free-ranging Formosan macaques usually submitted immediately after aggression (82-89%). The proportion of counter aggression in captive and free-ranging Formosan macaques were relative low (9-16%). The aqerage conciliatory tendency for adult Formosan macaques was 14.3% to 19.6%. The affiliative contacts in PC and MC in captive and free-ranging Formosan macaques were striking that both preferred grooming. The Formosan macaques significantly reconciled more during PC than MC period both in captive and free-ranging conditions. In addition, both had significantly more attracted than dispersed PC-MC pairs. The conciliatory tendencies in captive and free-ranging Formosan macaques were similar regardless of kin and non-kin partners. This study indicated that Formosan macaques were close to the macaques of Fascicularis group. Therefore, Formosan macaques had a despotic dominance style as suggested by Phylogenetic hypotheses.