黑长臂猿(Hylobates concolor)鸣叫行为研究

本文基于对云南黑长臂猿多年的野外研究,报道其鸣叫行为的一些特性。黑长臂猿通常在上午鸣叫,且大多数发生在930分以前,鸣叫的起始时间具季节性变化,有两个较为集中的时间,一个在730分左右,另一个在830分左右,然而起始时间并不与日出时间一致。黑长臂猿平均每两天鸣叫一次,日鸣叫发生频次在50％左右,鸣叫的发生也随着季节而变化。鸣叫的持续时间无群体的差异,一次鸣叫的平均时间在12分钟左右（除了GG群）,但群体在不同季节鸣叫时间长短不一。本文的研究结果同时还表明一个群体的鸣叫并未引起邻近其它群体的鸣叫。     

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.     

Singing by male and female Kloss gibbons (Hylobates klossii) in the Peleonan Forest, Siberut Island, Indonesia

Kloss gibbons (Hylobates klossii) are endemic to the Mentawai Islands in Indonesia and are one of only two gibbon species in which mated pairs do not sing duets. This is the first long-term study of the factors influencing the singing activity of Kloss gibbons within a northern Siberut Island population and follows two previous studies in central Siberut nearly 30 years ago. We collected data on the presence/absence of male and female singing within the study area on 198 days and within a focal group on 47 days. Rainfall during the time period in which they normally sing inhibits singing in both males and females. Our study supports the hypothesis that male and female songs function in intrasexual resource defence, as singing is associated with singing by same-sex neighbours, and same-sex choruses are more likely to occur after one or more days of silence (from that sex), suggesting there is pressure for individuals to communicate with same-sex neighbours regularly. Singing was not coordinated within a mated pair, suggesting that vocal coordination of the pair has been lost with the loss of the duet and that Kloss gibbon songs do not convey information to neighbours about the strength of the pair bond. On days when males sang predawn, females were more likely to sing after dawn and earlier in the morning. Additionally, the number of groups singing in female choruses was positively associated with the number of males that had sung in the predawn male chorus. We suggest that female songs have an intersexual territory defence as well as an intrasexual function.     

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.     

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.     

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.     

黑长臂猿的分布、现状与保护

黑长臂猿 (Hylobates concolor)是现生 1 1种长臂猿中其系统地位最低的灵长类。历史时期长臂猿曾广泛分布在我国南部的大部省区。自公元 4世纪以来 ,它们的分布发生了很大变化。分布区从北到南 ,从东到西急剧缩小 ,现黑长臂猿缩小到只在海南岛、云南南部和越南北部 ,已分化为 6个亚种。分布于我国的 5亚种中 ,海南亚种 (H . c. hainanus)现今仅20余只 ,是最濒危的一个亚种 ;.北部湾亚种 (H.c.nasutus) 50年代曾在广西西南部发现 ,60 -70年代已绝迹 ;指名亚种 (H.c.concolor)主要分布于越南北部、滇南和滇中哀牢山。滇南约有100余只 ,滇中哀牢山可能有40 - 60群 ,180 - 240只 ;景东亚种 (H . c. jingdongensis)为滇中无量山的特有亚种 ,现有100 - 116群 ,430 - 500只 ;滇西亚种 (H . c. furvogaster) ,只分布在滇西南的沧源、镇康、云县和耿马等地 ,约有 2 6-42群 ,100 - 150只。现今 ,黑长臂猿的分布区已不足1000Km2 ,总数量约1000只 ,为高度濒危的灵长类动物。造成黑长臂猿濒危的主要原因是 :热带和南亚热带原始森林的被破坏和缩小、人类活动的干扰使生境破碎和恶化、过度猎捕和长臂猿自身的生物学弱点。目前 ,中国的黑长臂猿已在 9个自然保护区中得到保护。     

东黑冠长臂猿鸣叫声谱分析

2007 年9 月至2009 年6 月,利用Sony TC - D5 Pro2 录音机、Sony C - 76 指向性话筒和Sony 录音磁带对广西邦亮自然保护区3 群东黑冠长臂猿的鸣叫进行了录音,对每群录音效果较好的5 个声音用Signal/ RTS 4.0 软件进行声谱分析。结果表明:东黑冠长臂猿叫声的频率较高,最高频率已经达到甚至超过了5 kHz;雄性的鸣叫声由起始音(boom)、简单的重复音节(aa notes)、调节前音节(pre-modulated note)和调节音句(modulated phrases)组成,雌性长臂猿一般只会发出一种固定而刻板的激动鸣叫,二者互相配合组成结构复杂的二重唱。通常,二重唱由成年雄性发起和结束,并占主导地位。同时,将东黑冠长臂猿二重唱的声谱结构与近缘种西黑冠长臂猿和海南长臂猿进行比较,结果显示雄性和雌性的叫声、以及二者配合发出的激动鸣叫序列在三者之间都有明显差异。因此,从声谱特征角度可以有效论证这三种长臂猿独立物种的分类地位。     

Preliminary observations of wild black-crested gibbons (Hylobates concolor concolor) in Yunnan province,people's republic of China

Black-crested gibbons (Hylobates concolor concolor), which inhabit the subtropical forests of southern China and northern Vietnam, have until now never been the subject of any systematic behavioral or ecological field study. This report presents the findings of a three-month field study of wild black-crested gibbons in the Wuliang and Ailao Mountain Game Reserves in Yunnan Province, People's Republic of China. The sites visited consisted of temperate, subtropical, broadleaf evergreen forests, with trees belonging to the families Elaeocarpaceae, Fagaceae, Lauraceae, Magnoliaceae, and Theaceae. Black-crested gibbons were documented from four sites visited and were found to be polygynous, with an average group size of seven to eight animals. The family groups were observed to comprise one adult male, from one to four adult females, and numerous offspring of various ages. Both sexes of this species emit interactive songs, and various features of this loud vocal behavior were analyzed and are discussed here. Since all other gibbon species were found to inhabit only tropical rain forests and live in strictly monogamous family groups, some of the ecological and evolutionary implications of these findings are also presented.     

黑长臂猿（Hylobates concolor）的配偶制及其与行为、生态和进化的关系

黑长臂猿是猿类中野外行为生态研究最少的一种,因而对其社群结构及配偶体制一直有着很大争议．本文基于近两年在无量山和哀牢山１０个月的野外工作,着重阐述了黑长臂猿的配偶体制,我们认为黑长臂猿是以一夫一妻和一夫多妻（至少是一夫两妻）的形式存在,并与其行为、生态及系统发育相关联．黑长臂猿的系统发育地位、成年雌性的领域性及群体的领域性不强和群体的领域较其它长臂猿类大得多可能是形成一夫多委的原因;而黑长臂猿在中国各分。布区的生境均受到不同程度的破坏,这在一定程度上也影响着其社群结构,特别是狩猎有着更直接的影响,活动范围的限制等,可能是形成一夫一妻的原因．     

Preliminary survey on the population and distribution of gibbons in yunnan province

Of the gibbons in China,Hylobates hoolock, H. lar, andH. concolor leucogenys are distributed only in Yunnan Province, whileH. c. concolor occurs limitedly in Yunnan Province and Hainan Island. During the periods from February to November 1984, and March to August 1985, the authors undertook surveys of the population and distribution of gibbons in 56 counties of 10 prefectures in Yunnan Province, and obtained the first reliable population and distribution data on the gibbons of this Province. The findings serve to fill an important blank in gibbon research in Yunnan and should pave the way for further research in the future.     

Singing behavior and singing functions of black‐crested gibbons (Nomascus concolor jingdongensis) at Mt. Wuliang,central Yunnan,China

We used data on loud duetted and solo songs collected from one habituated polygynous group of black‐crested gibbons (Nomascus concolor jingdongensis) on Mt. Wuliang, Yunnan, to test several hypotheses about the functions of these songs. The major functions proposed for loud gibbon songs include resource defense, mate defense, pairbonding, group cohesion and mate attraction. Duet bouts are generally initiated by adult males, who select the highest trees near to ridges or on steep slopes as singing trees. Such trees facilitate voice transmission and inter‐group communication. Singing trees tended to be located near important food patches and sleeping sites, which supports the resource defense hypothesis. The adult male and two adult females always sang interactively, alternating male phrases with the females' stereotyped great calls, to produce the duets, and females rarely produced great calls if they were more than 30 m from the male. The two females usually produced great calls synchronously during the duet, especially when they were close together. These features support both the mate defense and pairbonding hypotheses. The number of great calls and their degree of synchrony transmit information about spatial relationships and possibly pairbond strength to members to neighboring groups and floating animals. During or after the duet bouts, the adult females and juvenile moved toward to the adult male; and group members maintained a close spatial relationship, which supports the group cohesion hypothesis. Other incidents observed suggest a mate competition role for duets. The adult male always sang when the females started duetting with the subadult male. The subadult male sang solo bouts, but they were not more frequent or longer than bouts initiated by the adult male. Although mate attraction is the likely function of subadult solos, it was not convincingly demonstrated. In conclusion, all hypotheses concerning the function of singing are supported by at least some of the data, and none can be excluded. Am. J. Primatol. 71:539–547, 2009. © 2009 Wiley‐Liss, Inc.     

黑长臂猿（Hylobates concolor）对人类和非人类捕食者的回避行为

捕食和避免捕在动物生存和进化过程中起着重要作用,本文报道了我们自１９９０年３月至１９９２年元月观察黑长臂猿对人类和非人类捕食者的一系列反映,在遇到人类（观察者）后,据离观察者的远近和受惊程度不同,它们表现出５种回避方式,而在遇到非人类捕食者时,成年雌雄性一起将捕食者引开,以保护其后代个体免遭捕食。文中还对这些不同的行为方式进行了讨论。成年雄性是群体的主要保护者,成年雌性在群体保护中也起到重要的作用     

Biology and behaviour of reintroduced gibbons

We observed and recorded the behaviours of gibbons undergoing rehabilitation, before and after release, to document the behavioural and social changes of gibbons in the rehabilitation program and develop criteria for determining the suitability of a pair of gibbons for release. Hylobates albibarbis were observed at the Kalaweit Gibbon Rehabilitation Project in Kalimantan, Indonesia. Data were collected on animals both pre- and post-release and on wild gibbons for comparison. Data presented here show that reintroduced gibbons are capable of surviving without human intervention. In addition, their behaviour is similar to that of wild gibbons in terms of activity budgets, position in the canopy, body posture, pair association (PA) and diet. Prior to this study, no attempt has been made to quantify the rehabilitation process for gibbons, and rehabilitation project personnel require data reporting all aspects of a release so that improvements can be made. It is important to report these data for the benefit of future releases. Criteria, based on the behaviour of wild gibbons, are proposed to assist rehabilitation centers in assessing the suitability of gibbon pairs for release.     

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.     

Reassessment of age of sexual maturity in gibbons (hylobates spp.)

From studies of both wild and captive animals, gibbons are thought to reach sexual maturity at about 6 to 8 years of age, and the siamang (Hylobates syndactylus) at about 8 to 9 years. However, a review of the literature reveals that in most cases the exact age of the maturing animals was not known and had to be estimated. This study presents seven case reports on captive gibbons of known age. Captive males of the white-cheeked crested gibbon (H. leucogenys leucogenys) and of the siamang (H. syndactylus) can breed at the age of 4 and 4.3 years, respectively. Similarly, hybrid females (H. lar × H. moloch) and siamang females can breed at 5.1 and 5.2 years, respectively. This finding may help to improve the breeding success of captive gibbon populations. It is not clear whether gibbons reach sexual maturity earlier in captivity or whether sexual maturity is also reached by 5 years of age in the wild. Possible implications for the interpretation of group size regulation and of reproductive strategies of wild gibbons are discussed.     

海南长臂猿栖息地结构分析

海南长臂猿主要栖息在热带沟谷雨林和山地雨林的高树上,对别种林型颇难适应。沟谷雨林和山地雨林植物群落垂直结构明显,乔木层次在3层以上,富巨型木质藤本和附生、寄生植物,为猿类创造了良好的食物和鸣寝条件。猿群喜欢在山体中部和中下部沟谷、坡壁林缘地段活动,罕去高海拔山巅或山脊林区。若避免过甚的人为干扰,在海南岛现有自然林条件下,猿群有可能得到发展。     

Population and distribution of western black crested gibbon (Nomascus concolor) at Ailao Mountain, Xinping, Yunnan

The western black crested gibbon (Nomascus concolor) is mainly distributed in Yunnan, China. Ailao Mountain is located in central Yunnan and divided into three prefectures and six counties. This mountain forms the principle distribution range for western black crested gibbon; however, there are no published data on the gibbon population inhabiting the Xinping administrative. Take the interview results conducted in 2007 and 2009 with local people as the reference, this study conducted an extensive field survey covering all possible habitats from November 2009 to January 2010 using call surveys. Among the one hundred and twenty-four gibbon groups which were confirmed across the Ailao Mountain, the largest known population of western black crested gibbons yet, 85 groups inhabit the national nature reserve and adjacent national forest, 30 groups inhabit the provincial nature reserve and nine groups inhabit the collective forest located outside the reserve and national forest. We found that the western black crested gibbons here have a patchy distribution pattern and occur at higher densities in certain areas. Moreover, the population distribution density and elevation gradient distribution decline from north to south. The results also demonstrated the importance of Ailao Mountain in the western black crested gibbon protection.     

A comparative study of crested gibbons (Nomascus)

Crested gibbons (Nomascus) are in the rarest genus of the family Hylobatidae, with the Hainan gibbon (Nomascus hainanus) being the rarest primate in the world. In the past, the number of species in this genus has been at the center of much controversy, in part, because their color changes during immaturity as well as other factors, such as physical similarities in genitalia, creating difficulties in accurately determining the sex of individuals. Furthermore, owing to their rarity, illusiveness, and the rough terrain that comprises their native habitat, Nomascus is one of the least studied Hylobatidae. This article represents the most comprehensive dissemination of visual characteristics of the genus Nomascus to assist in the accurate identification of captive and wild crested gibbons. Through differences in pelage color, skeletal anatomy, dentition, vocalizations, behavior, distribution, and genetic studies, we are able to determine more accurately whether or not a subspecies should be elevated to species level. From the current data, there are six species and one subspecies in the genus Nomascus. However, reports of a recently identified light-cheeked gibbon (Nomascus sp.) in northeast Cambodia, Central Vietnam, and South Lao PDR, will add additional taxa to this genus.     

滇南、滇东南黑冠长臂猿分布与数量

滇南、滇东南曾是黑冠长臂猿广泛分布的地区之一。为掌握该地区近年黑冠长臂猿分布及种群数量现状,从2003年7月~2004年12月,采用访问调查及利用鸣声定位法进行了实地调查。调查结果显示,在近40000km2的范围内,东黑冠长臂猿(Nomascussp.cf.nasutus)在云南境内可能已消失;西黑冠长臂猿(N.concolor)亦仅发现4~7个种群,且孤立分布于3个地区(金平芭蕉河2群6只,金平西隆山1-2群,绿春黄连山1~3群,总计不超过25只),而在江城牛倮河自然保护区、马关古林箐自然保护区、麻栗坡老君山自然保护区及屏边大围山自然保护区可能已绝迹。滇南、滇东南黑冠长臂猿分布区缩小及种群数量剧减,主要与栖息地丧失及过度捕猎有关。