The ecology of singing in Kloss gibbons (Hylobates klossii) on Siberut Island,Indonesia

Data are presented from a study of habituated Kloss gibbons on Siberut Island, Indonesia. Male Kloss gibbons can sing at any time from 0100 to 1300 hr, but the majority of songs is concentrated in the hour before dawn. Female Kloss gibbons sing only after dawn and the song bout includes a dramatic visual display. Neither countersinging nor coordinated chorusing has been proved in either sex. Males sing before dawn as often as possible but are inhibited by wet nights and by minimum temperatures below 21.5°C; postdawn songs of both sexes are inhibited by rain. The occurrence of any particular type of song bout is independent of the occurrence of the other types. Song trees used by males and those used by females do not differ in height. Song trees emerged from the neighboring canopy more than other available trees of similar height in the gibbons’ home range. Female song trees were most abundant on the slopes and where the trees were tallest. Almost all the male’s night trees could have been used for singing from had the weather been suitable. There was a greater likelihood of the male’s traveling a long way to the day’s first fruit source on mornings when he sang before dawn than on mornings when he did not. Considerations of sound transmission through tropical rain forest reveal that the times and frequencies used for singing by Kloss gibbons are optimal for communicating with neighboring groups.     

Calling in wild silvery gibbons (Hylobates moloch) in Java (Indonesia): behavior, phylogeny, and conservation

Hardly any behavioral data are available for the silvery gibbon (Hylobates moloch), an endangered primate that is endemic to the island of Java, Indonesia. We studied the singing behavior of the easternmost population of this species in the Dieng mountains, central Java, in 1998-1999. We aimed to document the timing of singing, quantify the amount of singing by the respective sexes, and explore the role of bioacoustics in density estimation. A total of 122 song bouts in at least 12 groups were monitored. No duet songs were heard. Most of the song bouts (91.5%) were female solo song bouts or female scream bouts. In contrast to an earlier study on the westernmost population of silvery gibbons, during which few if any male songs were heard, at least 8.5% of the song bouts in our study were male solo song bouts. They were significantly longer in duration than the female songs. All male song bouts uttered before dawn (0520 hr) were produced in a chorus fashion, with at least three individuals participating. Choruses occurred about once every 8.5 days, and lasted longer and occurred earlier than female solo song bouts. Most male songs (60%) started between 0355-0440 hr, when it was still dark. All female songs, in contrast, started after 0500 hr, and female singing activity peaked around 0600. Regular male singing, male chorusing, and regular predawn singing have not previously been reported for silvery gibbons. Similarly separated periods of male and female solo songs and the absence of duetting have been observed in Kloss's gibbons (H. klossii) on the Mentawai Islands, and may represent synapomorphies shared by both species. The pronounced individual-specific song characteristics of silvery gibbons allow accurate mapping of groups. The density of gibbons at our study site was established to be 1.9-3.7 groups/km2, corresponding to 6.7-13.1 individuals/km2. We reassess the suitability of gibbon songs as a means of estimating the density and size of gibbon populations, and discuss the proximate causes for the absence of duetting in silvery gibbons.     

Vocal responses of captive gibbon groups to a mate change in a pair of white-cheeked gibbons (Nomascus leucogenys)

The singing behaviour of 3 pairs of white-cheeked gibbons (Nomascus leucogenys) held at the Perth Zoo was observed for 6 months in 2005. These groups included a family (mated pair and 2 immature offspring) and a pair without offspring. During the study, the female without offspring was exchanged for an unpaired female from New Zealand. After the new pair had been released onto the island enclosure and began to duet, the duetting rate of the white-cheeked gibbon family increased. The increased singing began after the new female had started to sing solo female great calls. These observations support the hypothesis that duets have an intergroup communication function in white-cheeked gibbons. The pair that duetted most frequently also copulated most frequently but allogroomed the least. We suggest that duetting may be more important to intergroup relations than to pair bond maintenance in this species.     

Sound Spectrum Characteristics of Songs of Hainan Gibbon (Nomascus hainanus)

Gibbons are characterized by their species-specific calls, or songs. There are few studies of songs of Hainan gibbon (Nomascus hainanus). To study the sound spectrum characteristics and test for intergroup differences in Hainan gibbon song, we studied the singing behavior of Hainan gibbons in Bawangling National Nature Reserve, Hainan Province, China, intermittently from August 2002 to February 2013, collecting 184 recordings. Our results show that: 1) Hainan gibbon song bouts occur mainly 0–4 h after dawn. 2) The songs of adult males living in groups are composed mainly of one to three short notes and one to five long notes, while solitary adult male songs consist only of long frequency modulated notes and no short or single notes. 3) The song chorus is dominated by adult males, while females add a great call. Males do not have a great call, unlike those in other gibbon species. There are no female solos. 4) The sound spectrum frequency is similar in adult males living in two different groups, but the duration of the first long note differed significantly between the groups. The sonic frequencies of male and female songs are lower than those of other gibbons: no more than 2 kHz. Hainan gibbon sound structure is simple, although females participate in the chorus, reflecting their primitive status among gibbon species.     

Female Songs of the Nonduetting Javan Gibbons (<Emphasis Type="Italic">Hylobates moloch</Emphasis>) Function for Territorial Defense

Duets in territorial, pair-living primates may function to maintain intragroup cohesion, promote intergroup avoidance, and assist in territorial and resource defense, as well as advertising and reinforcing pair bonds. Despite the absence of duetting in Javan gibbons (Hylobates moloch), recent playback experiments suggested that Javan gibbon songs also play a role in pair-bond advertisement as well as territorial and mate defense. However, playback experiments only assess motivations of the listener, which may not reflect the motivations of the caller. We conducted an observational study of naturally occurring female songs in two groups of Javan gibbons from July 2009 to March 2010 and from March to November 2011 in Gunung Halimun–Salak National Park, Indonesia. We investigated female singing rates in relation to singing location, daily path length, occurrence of intergroup encounters, feeding rate, allogrooming rate, and distance between pair mates. The two females produced 47 songs during 164 observation days. Females in the area of their home range that overlapped with neighboring groups sang more frequently than expected based on time spent in the area of overlap vs. the home range interior. Groups also had longer daily path lengths on days when females sang than on nonsinging days, and on days when they visited the area of overlap than on nonvisiting days. Our findings indicate that, like the duets of other pair-living territorial primates, female Javan gibbon songs function for territorial defense, but we found no support for other functions such as intergroup avoidance, resource defense, and pair-bond reinforcement.     

Duet-splitting and the evolution of gibbon songs

Unlike the great apes and most other primates, all species of gibbons are known to produce elaborate, species-specific and sex-specific patterns of vocalisation usually referred to as "songs". In most, but not all, species, mated pairs may characteristically combine their songs in a relatively rigid pattern to produce coordinated duet songs. Previous studies disagree on whether duetting or the absence of duetting represented the primitive condition in gibbons. The present study compares singing behaviour in all gibbon species. Various vocal characteristics were subjected to a phylogenetic analysis using previously published phylogenetic trees of the gibbon radiation as a framework. Variables included the degree of sex-specificity of the vocal repertoire, the occurrence of solo songs, and the preference for a specific time of day for song-production. The results suggest the following scenario for the evolution of gibbon songs: (1) The last common ancestor of recent gibbons produced duet songs. (2) Gibbon duets probably evolved from a song which was common to both sexes and which only later became separated into male-specific and female-specific parts (song-splitting theory). (3) A process tentatively called "duet-splitting" is suggested to have led secondarily from a duetting species to a non-duetting species, in that the contributions of the pair-partners split into temporally segregated solo songs. This appears to be the first time that a non-duetting animal can be shown to be derived from a duetting form. (4) The return to exclusive solo singing may be related to the isolated island distribution of the non-duetting species.     

Individuality in the songs of wild agile gibbons (Hylobates agilis) of Peninsular Malaysia

Recordings were made and analyzed of the female dominated duet songs by eight adult pairs of wild agile gibbons from two sites in Peninsular Malaysia. A statistical analysis of their songs revealed that individuality occurred throughout the females' songs from both sites. Individuality in gibbon songs may allow singing individuals to more efficiently and effectively locate and identify neighboring nonmate conspecifics.     

The formation of a new gibbon group

Families of gibbons and siamang were studied in lowland dipterocarp forest at Kuala Lompat, Pahang, Malaysia between January and July 1973. In April a lone male gibbon (who had previously been a frequent associate of a lone male siamang) attracted a female gibbon and the two animals called together in chorus. The new pair were immediately attacked and separated by a long-established neighbouring family of gibbons. Four days later the new pair reformed and despite further fierce attacks from their neighbours remained together and gradually established their own territory. The lone siamang also attracted a female into his range. The new siamang pair stayed together for most of four days but eventually the female left his range and did not return. Generalisations are made concerning the way that new hylobatid groups become establilished.     

Do female great tits (Parus major) assess males by eavesdropping? A field study using interactive song playback

Male singing behaviour correlates with extra-pair success in several passerine birds. Singing interactions during territorial contests provide relative information on the males involved. Such information may be important in female extra-pair behaviour and eavesdropping on singing interactions among males may allow females to make such relative assessments. We used interactive playback to instigate singing contests with male great tits during the peak fertile period of their mate in an attempt to alter females'' assessment of mates'' quality relative to neighbours (potential extra-pair partners). We escalated a contest to one male (by overlapping his songs) and then subsequently de-escalated a contest (by alternating) to a neighbour. Intrusions onto neighbouring territories by females mated to either treatment male were then monitored. Females mated to escalation treatment males were more likely to intrude following playbacks than females mated to de-escalation treatment males. Although the absolute song output of males did not differ between treatments, males produced more song relative to playback in de-escalation treatments and relative song output was positively correlated with female intrusions. Therefore, female great tits eavesdrop on singing interactions and change their visitation rates to neighbouring territories according to their mate''s singing performance relative to neighbours.     

Experimental field studies of Asian ape social systems

The Asian apes, orangutans and gibbons, possess unusual social systems among anthropoid primates. Social groups of gibbons consist of mated adult pairs and their offspring; mature orangutans are primarily solitary. Recent experimental field research has begun to yield insights into the behavioral mechanisms employed by these animals to maintain their characteristic patterns of social dispersion. While spatial separation between female orangutans appears to be maintained passively, aggression, which is manifest during direct encounters and long-distance vocal interactions, mediates male asociality. Male-male aggression is the result of intense intrasexual competition occurring between animals for mating access to females. To reduce intrasexual competition, male orangutans have adopted alternative mating tactics. In contrast to female orangutans, female gibbons show marked agonistic tendencies toward conspecifics. Female territoriality contributes to preventing males from becoming polygynous. Male gibbons, restricted to monogamous relationships, attempt to ensure their paternity through intrasexual aggression. These observations suggest that the spatial dispersion of females constrains male mating options in both species. However, variations between orangutan and gibbon social systems can be understood as consequences of the temporal dispersion of sexually receptive females. The temporal clumping of females, due to relatively high operational sex ratios, limits the ability of male gibbons to acquire multiple mates. Conversely, an extremely low operational sex ratio in orangutans creates a strong selection pressure for intrasexual competition and polygamous mating. These considerations provide a novel framework for interpreting the social systems of the African apes.     

Flexibility in the species-typical songs of gibbons

Flexibility, or a capacity for situational variation, is an important characteristic of gibbon vocalization—one which may be seen as necessary to the functional effectiveness of species-typical songs. This paper reviews literature reporting flexibility of gibbon vocal behavior in relation to reinforcement contingencies, the singing of neighboring gibbons, development of pair coordination in the duet-singing of siamang gibbons, sequential progression in the elaboration of organizing sequences in siamang gibbons, and “repairs” of organizing and great-call sequences. A theoretical framework to account for the development of flexibility in species-typical behaviors is drawn on the basis ofGlickman's andSchiff's (1967) proposal of the reinforcing power of species-typical behaviors and associated feedback and consequent stimuli.     

Territory and monogamy among Kloss' gibbons (Hylobates klossii) in Siberut Island, Indonesia.

Behavior of Kloss' gibbons was studied from July 1 to October 7, 1972 in Siberut Island, off the coast of western Sumatra, Indonesia. Reproductive groups are monogamous families with a mean family size of 3.4 individuals (n = 11 families). Such families occupy territories averaging 6.7 ha. Adults defend their territories only against members of the same sex. This intrasexual defense of territory maintains the monogamous mating system. Other social units were unmated resident females, unmated resident males, floating males, and a courting pair. Males establish territories before mating, perhaps with help from their parents. Males guard their families against predators. Females lead progression through the territory. Subadults remain peripheral to their families but other family members tend to remain within 10 m of one another.     

Home range use by Kloss gibbons (Hylobates klossii) on Siberut Island,Indonesia

A home range is a reservoir of resources which are distributed throughout its area and joined by pathways. These resources are used for different activities and at different times of day and while ‘space-time systems’ have been described for a few terrestrial mammals, no previous study has attempted a similar analysis for an arboreal mammal. Kloss gibbons were studied for two years on Siberut Island, Indonesia, and one group was habituated to an observer. The ranging patterns of this group are analysed in two, three and four dimensions with respect to the types of forest in which the animals lived. Finally, differences between the results of this and previous studies of Kloss gibbons are discussed and it is suggested that a home range can be more permanent than the animals within it.     

Song dialects,mate selection,and breeding success in white-crowned sparrows

The songs of 15 colour-marked, mated pairs of white-crowned sparrows (Zonotrichia leucophrys nuttalli) were recorded in a contact zone between the Presidio and San Francisco, California dialect regions. Males were recorded in the field and females were trapped after the breeding season and injected with testosterone to induce singing. The song of each member of a mated pair was classified using two different markers. It was found that most pairs did not match song types, and that the reproductive success of those that matched did not differ from those that did not match. Most of the territorial males sang the Presidio dialect, while most injected females sang the San Francisco dialect. The results fail to support a positive assortative mating hypothesis regarding the functional significance of learning dialects. It is suggested that the mismatching of songs could be due to sexual differences in the disposition to learn songs of neighbours at the breeding site.     

The organization of song in the Hainan black gibbon (Hylobates concolor hainanus)

There have never been any documented studies on the vocal or non-vocal behaviour of wild or captive Hainan black gibbons, which are highly endangered in the wild (Hainan Island, China), and very scarce in captivity. The singing behaviour of the only pair of captive Hainan black gibbons outside Southeast Asia was observed, recorded and analyzed in detail. This adult pair produced duets only, with the organizational features of their duets found here to be similar to that in two other concolor gibbon sub-species; however, the acoustical features were very different from any other concolor gibbon pair studied previously. Some functional and evolutionary implications of the singing behaviour of the Hainan black gibbon are presented and discussed.     

Function of pair duets in the eastern whipbird: cooperative defense or sexual conflict?

Paired male and female eastern whipbirds, Psophodes olivaceus,sing precisely coordinated, male-led duets. Four broad explanationshave been proposed for the function of duets: 1) cooperativeresource defense, 2) prevention of partner usurpation, 3) defenseof an individual's own position within the partnership, or 4)mate identification and localization. These 4 hypotheses makedifferent predictions about how male and female residents shouldrespond to simulated intrusion by other pairs or individuals.We compared the behavioral and vocal responses of 20 pairs ofeastern whipbirds to simulated territorial intrusions by: 1)a solitary singing male, 2) a solitary singing female, and 3)a duetting pair. Males and females did not coordinate theirapproach to the playback speaker and showed sex-specific responsesto playback. Males did not respond differently to duetting versussolo singing intruders. By contrast, females approached moreclosely during solo female song than during solo male song orduet playback. Females also produced specific vocalizationsonly in response to duet and solo female playback. Both sexesapproached the speaker more closely and quickly during playbackof same-sex solo songs than opposite-sex solo songs. Finally,females answered more of their mate's songs during simulatedintrusion by a lone female than during simulated intrusion bya lone male. Our results suggest that duets in this speciesprimarily function to allow females to defend their exclusiveposition in a partnership. Mate defense by females is unusualin birds but may be promoted in eastern whipbirds by a female-biasedsex ratio and the need for exclusive access to male care. Thus,duets result from independent and conflicting strategies ofmate and territory defense in males and females.     

Social factors affect the singing rates of female northern cardinals Cardinalis cardinalis

Social factors, such as the duration of territorial occupancy or of time paired with a mate can affect the rate at which individual birds sing. This study examined the influence of duration of pair‐bond and territory occupancy as well as date on the rate of singing by male and female northern cardinals Cardinalis cardinalis. When differences in breeding status were controlled, female cardinals sang at higher rates earlier in the season. Females in newly formed pairs sang at significantly higher rates than those in pairs that had previously bred together. In contrast, male cardinals did not show significant variation in the rate of singing throughout the season. The song rates of males in newly formed and established pairs did not differ significantly. Song rates for males and females in mated pairs were not significantly correlated. This study suggests that social factors have a strong effect on the rate at which female cardinals sing. It is possible that increased intra‐sexual aggression by females when they are establishing a new territory with a new mate leads to this higher level of song output. Once females have established a territory and acquired a mate, dear‐enemy effects might diminish the need for acoustic territorial defence. Differences in the constraints of nesting or the attraction of extra‐pair mates might explain the sexual differences in male and female singing behaviour.     

Song Functions in Nonduetting Gibbons: Evidence from Playback Experiments on Javan Gibbons (<Emphasis Type="Italic">Hylobates moloch</Emphasis>)

Territorial, pair-living primates usually perform long-distance calls as duets in which adult males and females coordinate their calls. Previous studies using playback experiments have shown that gibbon duets convey information about the status of the caller (location, familiarity, sex of the caller, and paired status) and gibbons use this information to respond to achieve several nonmutually exclusive functions, including intragroup contact, territorial defense, and pair-bond advertisement and strengthening. However, not all pair-living gibbons duet, and it is unclear whether the same results should be expected in nonduetting species. We conducted song playback experiments (N = 47 trials) to test hypotheses about song functions in nonduetting gibbons on two groups of wild Javan gibbons (Hylobates moloch) in the Gunung Halimun-Salak National Park, Indonesia. Javan gibbons initiated movement toward the speaker more quickly in response to songs broadcast in the center of the territory, stranger songs, and songs of unpaired individuals than to songs at the border, neighbor songs, and songs from paired individuals. These results suggest that Javan gibbons can localize songs, and that Javan gibbon songs transmit information about the identity and paired status of the caller. Our results imply that Javan gibbon solo songs are likely to function for territorial defense and pair-bond advertisement like duets in other primates.     

Immature male gibbons produce female-specific songs

Gibbons are apes that are well known to produce characteristic species-specific loud calls, referred to as “songs.” Of particular interest is the sex specificity of the “great calls” heard in gibbon songs. However, little is known about the development of such calls. While great calls are given by female gibbons of various ages, they have never been recorded from males. Here, we report two observations of immature male gibbons from two different species, wild Hylobates agilis and captive H. lar, which spontaneously sang female-specific great calls. Based on the video clips, we conclude that immature males also have the potential to produce great calls. Our observations led us to propose a new hypothesis for the development of sexual differentiation in the songs of gibbons, and its implications for the general issue of sex-specific behavior in primates.     

云南高黎贡山大塘东部白眉长臂猿鸣叫行为

通过2009年10月至2010年7月监测高黎贡山大塘的3群东部白眉长臂猿的鸣叫行为,收集了10个月(200d)的数据,发现东部白眉长臂猿的鸣叫行为主要发生在黎明后4h内;平均每次鸣叫持续14min59s,不同群体间鸣叫持续时间不同。天气情况影响东部白眉长臂猿的鸣叫开始时间(相对黎明)和持续时间,在雨天鸣叫相对黎明开始时间较晚,持续时间也较长。东部白眉长臂猿鸣叫频率在10月和11月明显高于其他月份,可能与食物(果实)的丰富度有关。根据研究结果,建议对东部白眉长臂猿的调查最好集中在10月或11月;每天监测时间至少持续至黎明后4h,以覆盖长臂猿90%以上的鸣叫;如果出现下雨天气,应适当延长调查时间。