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.     

Status and distribution pattern of black crested gibbon (Nomascus concolor jingdongensis) in Wuliang Mountains, Yunnan, China: implication for conservation

The western black crested gibbon (Nomascus concolor), or black gibbon, one of the lesser apes (Hylobatidae), is mainly distributed in Yunnan, China. Of the four recognized subspecies, N. c. jingdongensis is endemic to the Wuliang Mountain, central Yunnan, China. Of all the subspecies, this one has the largest population of any black gibbon. However, the data were all based on brief estimates. We carried out an extensive field survey on population and group distribution of the black gibbon in the Wuliang Mountains by use of loud morning songs and interviews with local people. Ninety-eight groups were confirmed and located in the mid-montane range of Wuliang Mountains. More groups are found on the east slope and the southern region than in the west and the north. Gibbons are more disjunctly distributed on the west slope, especially in the northern part. Deforestation in the late 1950s, 1960s and 1970s was the main reason for rapid loss of habitat and population decline. Hunting was another key reason for population decline and, in many cases, the main reason for local extinction.     

Predation on giant flying squirrels (Petaurista philippensis) by black crested gibbons (Nomascus concolor jingdongensis) at Mt. Wuliang, Yunnan, China

Predation on vertebrates is infrequent in gibbons. In a 14-month field study of the central Yunnan black crested gibbon (Nomascus concolor jingdongensis) at Mt. Wuliang, Yunnan, China, we observed gibbons attacking, killing and eating giant flying squirrels (Petaurista philippensis). During 845 h of observation on one study group, the gibbons attacked giant flying squirrels 11 times, and succeeded in 4 cases. Although all members of the group attempted to attack the squirrels, all four successful attacks were made by the same adult female. The victims were infants in three cases and a juvenile or sub-adult in one case. Black crested gibbons also attacked adult giant flying squirrels by grabbing their long tails and throwing them from the canopy, but they failed to catch or kill the prey in three cases observed. Passive meat sharing occurred in three out of four successful cases. Besides hunting giant flying squirrels, the black crested gibbons also ate eggs or chicks in two birds’ nests and one lizard.     

Influence of food availability and climate on behavior patterns of western black crested gibbons (Nomascus concolor) at Mt. Wuliang,Yunnan, China

Food abundance and climatic factors can significantly affect the behavior of animals and constrain their activity budgets. The population of western black crested gibbons (Nomascus concolor) in Mt. Wuliang lives in montane forest and is close to the northern extreme of the distribution for gibbons (Hylobatidae). Their habitats show remarkable seasonal variation in terms of food availability, temperature, and rainfall. To understand behavioral adaptations of western black crested gibbons to different sets of ecological conditions, we examined relationships among food availability, mean temperature, rainfall, and behavior patterns by observing two groups for 1 year each. Our results revealed that activity budget was affected by food availability and mean temperature. The gibbons spent more time eating flowers when that resource was more available and spent less time moving when fruit was more available. The gibbons spent less time feeding and more time resting, and spent less feeding time on fruit and leaves when the mean temperature was lower. These results suggest that the gibbons displayed a pronounced preference for flowers as a food resource and adopted a time minimizer strategy when high‐nutrient food items (i.e., fruit) were more available. In addition, the gibbons adopted an energy‐conserving strategy during periods of low temperature. The flexibility of behavioral patterns in responding to food availability and temperature may potentially improve the gibbons’ prospects of surviving and reproducing in a northern montane forest.     

无量山大寨子黑长臂猿(Nomascus concolor jingdongensis)种群生存力

据2003年9月至2005年9月,对云南中部无量山大寨子黑长臂猿种群(5个群体)进行了观察,获得了群体大小、配偶体制、繁殖间隔、环境容纳量、死亡率、灾害的发生频率等种群参数,并结合近缘种的一些相关数据,利用旋涡模型(Vortex 914),对无量山大寨子地区黑长臂猿亚种群的动态进行了模拟分析。结果显示:大寨子亚种群是一个具有很强的潜在繁殖力的种群,如果没有偷猎,亚种群在100a之内不会灭绝,并且能迅速达到环境容纳量。但是每年如果有1只成年雄性和1只成年雌性被猎杀,该种群将会在第78年灭绝,且灭绝概率为100%。不同程度的死亡率对种群影响不大,但高死亡率显著延缓了种群到达环境容纳量的时间。环境容纳量对种群遗传多样性损失具有重要的影响,在没有猎杀的情况下,种群的长期存活需要一个较大的环境容纳量。因此,在黑长臂猿受到严格保护、且栖息地主要在保护区内的今天,严密监控火灾的发生,限制牲畜进入林区等人为干扰的影响,保护好黑长臂猿栖息地是首要工作之一。但如果能使其栖息地周围的森林植被得到恢复,增加其栖息范围,将有利于该地区黑长臂猿的发展。     

无量山大寨子黑长臂猿一夫二妻制的群体结构及其行为学原因

2003年8月—2005年8月,对无量山大寨子5个黑长臂猿群体的结构和组成进行了观察。当一个群体在早晨鸣叫或依次通过树冠时,记录群体的结构和组成。每个群体都由1个成年雄性、2个成年雌性及其后代组成。2003年8月平均群体大小为6·2只;到2005年8月,平均群体大小发展为6·4只,其中有2个亚成年雄性从出生群迁出,且有3只幼猿出生。在3个群体(G1、G2和G3)中两个成年雌性都成功繁殖了后代。同一群体内两个成年雌性间无攻击或等级行为。2005年4月15日,当一只亚成年雌性进入G3的领域后,两只成年雌性对其进行追逐驱赶,并且干扰其与成年雄性配合进行二重唱,成年雄性没有直接驱赶流浪的亚成年雌性,10天后这只亚成年雌性离开了G3的领域。亚成年雄性经常与群体其他成员保持一定距离,并且在出生地通过独唱练习鸣叫。黑长臂猿可能通过亚成年雄性和雌性的迁出,及成年雌性对外来流浪雌性的驱赶维持这种一夫二妻的群体结构。     

Proximity association in polygynous western black crested gibbons (Nomascus concolor jingdongensis): network structure and seasonality

We investigated the structure and seasonality of the proximity network in a group of polygynous western black crested gibbons (Nomascus concolor) using social network analysis. The spatial proximity changed seasonally and was affected by temperature and rainfall. Preferred proximity association was not distributed randomly among individuals. Kinship was one explanation for the social structure, as offspring preferred to maintain close proximity with their mothers. The proximity of infants to mothers decreased with age, and independent offspring had lower proximity to mothers than dependent ones. We found that the adult male had different proximity relationships with two different adult females. The frequency of proximity between the male and the infant-carrying female was significantly higher than that between the male and the female who had immigrated carrying one offspring of uncertain paternity into the group. Infanticide avoidance and/or predation protection for dependent infants might explain the proximity relationship differences. Temperature influenced group proximity association, with individual proximity increasing in the cold months and decreasing in the hot months. Group proximity decreased in months with higher anthropogenic disturbance.     

云南中部无量山黑长臂猿(Nomascus concolor jingdongensis)栖息地乔木层物种多样性和结构特征

无量山是黑长臂猿景东亚种的唯一分布区,现栖息着98群500余只黑长臂猿。长臂猿是典型的树栖性灵长类动物,通过对无量山西坡大寨子地区250个样方的调查,分析了随海拔梯度升高,黑长臂猿栖息地植被乔木层物种多样性和结构的变化情况。共记录乔木111种,隶属于33科;木质藤本23种,隶属于18科。其中优势科主要有樟科(Lauraceae)、木兰科(Magnoliaceae)、壳斗科(Fagaceae)、山茶科(Theaceae)和杜鹃花科(Ericaceae)。根据不同海拔乔木层物种组成的差异,该区可划分为半湿润常绿阔叶林(2100~2300m)和中山湿性常绿阔叶林(2400~2600m)两种植被类型。多样性指数(Shannon-Wiener指数)随海拔升高在两种植被类型中均呈下降趋势;均匀度指数(Pielou指数)在半湿润常绿阔叶林中呈下降趋势,在中山湿性常绿阔叶林中无明显变化。半湿润常绿阔叶林中,阳坡的乔木物种多样性低于阴坡;而阳坡物种多样性在两种植被类型间差异显著。随着海拔的升高,乔木层的平均高度和上层乔木(Ⅰ层)的比例均逐渐降低,但下层乔木却呈上升趋势。6种优势乔木树种的种群年龄结构分析表明,该区植被的乔木层呈现稳定增长的趋势。     

云南无量山黑长臂猿对植物种子的传播作用

2005年9月至2006年4月,对无量山一群黑长臂猿进行观察,研究黑长臂猿在植物种子传播中的作用.观察记录黑长臂猿一天的排便次数并收集粪便.通过直接观察和收集长臂猿粪便确定长臂猿取食的果实种类.以森林中采集的果实种子为对照标本,对粪便中的种子进行鉴定,并记录各种种子的数量,测量其长度、宽度和重量.结果表明观察期间黑长臂猿共取食31种果实,其中有1种在被取食时尚未成熟,另有2种果实的种子在取食后遭到破坏,还有1种植物的种子不能被长臂猿吞食而得不到传播,黑长臂猿可为27种植物传播种子.黑长臂猿个体平均每天排便2次,粪便的平均湿重为22.7 g,每份粪便平均含有1种植物的种子;在不包含小种子(直径<3 mm)的情况下,平均每份粪便中含有12粒种子.总的来说,黑长臂猿是有效的种子传播者.     

西黑冠长臂猿雄性取代前后鸣叫行为的变化

本文通过长时间的野外监测,首次报道了西黑冠长臂猿的一次雄性取代行为。一个研究群体(G3)中的1 只亚成年雄性长臂猿在10 岁左右取代了相邻群体(G2)中的成年雄性。整个取代过程持续了15 d时间。雄性取代发生前,G2 中雌雄的配对关系已经不稳固,这为雄性取代提供了机会。而G2 与G3 群的一次长时间冲突可能消耗了G2 中成年雄性大量体能,这为G3 中的亚成年雄性打败并取代G2 中的成年雄性创造了机会。本研究在取代发生后,对新形成群体的鸣叫行为进行了连续4 个月的监测。结果表明与处于稳定时期的G2 群相比,新形成群体的鸣叫频次更高,但每次二重唱中雌性的平均激动鸣叫次数降低。这证明了Geissmann (1986)提出的假说,新配对的群体应该在尽量短的时间内多练习二重唱,这样导致新配对群体的鸣叫频率明显升高。虽然经历了4 个月的合唱练习,新形成群体的激动鸣叫次数仍然偏低,并且两只雌性同时激动鸣叫的频次也比较低。这说明新形成的配对之间配合依然不默契,或者说明配对之间的关系还不稳定。     

Phylogeny and distribution of crested gibbons (genus Nomascus) based on mitochondrial cytochrome b gene sequence data

Crested gibbons, genus Nomascus, are endemic to the Indochinese bioregion and occur only in Vietnam, Laos, Cambodia, and southern China. However, knowledge about the number of species to be recognized and their exact geographical distributions is still limited. To further elucidate the evolutionary history of crested gibbon species and to settle their distribution ranges, we analyzed the complete mitochondrial cytochrome b gene from 79 crested gibbon individuals from known locations. Based on our findings, crested gibbons should be classified into seven species. Within N. concolor, we recognize two subspecies, N. concolor concolor and N. concolor lu. Phylogenetic reconstructions indicate that the northernmost species, N. hainanus, N. nasutus, and N. concolor branched off first, suggesting that the genus originated in the north and successively migrated to the south. The most recent splits within Nomascus occurred between N. leucogenys and N. siki, and between Nomascus sp. and N. gabriellae. Based on our data, the currently postulated distributions of the latter four species have to be revised. Our study shows that molecular methods are a useful tool to elucidate phylogenetic relationships among crested gibbons and to determine species boundaries. Am. J. Primatol. 72:1047–1054, 2010. © 2010 Wiley‐Liss, Inc.     

无量山大寨子黑长臂猿(Nomascus concolor jingdongensis)种群生存力

据2003年9月至2005年9月,对云南中部无量山大寨子黑长臂猿种群（5个群体）进行了观察,获得了群体大小、配偶体制、繁殖间隔、环境容纳量、死亡率、灾害的发生频率等种群参数,并结合近缘种的一些相关数据,利用旋涡模型（Vortex 914）,对无量山大寨子地区黑长臂猿亚种群的动态进行了模拟分析。结果显示：大寨子亚种群是一个具有很强的潜在繁殖力的种群,如果没有偷猎,亚种群在100a之内不会灭绝,并且能迅速达到环境容纳量。但是每年如果有1只成年雄性和1只成年雌性被猎杀,该种群将会在第78年灭绝,且灭绝概率为100%。不同程度的死亡率对种群影响不大,但高死亡率显著延缓了种群到达环境容纳量的时间。环境容纳量对种群遗传多样性损失具有重要的影响,在没有猎杀的情况下,种群的长期存活需要一个较大的环境容纳量。因此,在黑长臂猿受到严格保护、且栖息地主要在保护区内的今天,严密监控火灾的发生,限制牲畜进入林区等人为干扰的影响,保护好黑长臂猿栖息地是首要工作之一。但如果能使其栖息地周围的森林植被得到恢复,增加其栖息范围,将有利于该地区黑长臂猿的发展。     

Population differences and acoustic stability in male songs of wild western black crested gibbons (Nomascus concolor) in Mt. Wuliang, Yunnan

Population differences and acoustic stability in male songs of the western black crested gibbon (Nomascus concolor) at Mt. Wuliang, Yunnan, were investigated using data collected over 4 years. Detectable vocal differences were found between the western and eastern populations within Mt. Wuliang. Discriminant functions calculated using the phrases recorded in 2007-2008 correctly classified phrases recorded in 2009, and discriminant functions produced from phrases recorded in 2007-2009 could correctly classify phrases recorded in 2010 for 4 of 5 males. Results showed that the male songs of western black crested gibbons remain relatively stable over years. In light of the acoustic individuality in male songs of this gibbon species, which has recently been proved, and the acoustic stability shown in this research, we suggest that the song of male individuals may be useful in monitoring individuals or family groups of this species.     

Seasonal Variations in the Activity Budget of <Emphasis Type="Italic">Nomascus concolor jingdongensis</Emphasis> at Mt. Wuliang,Central Yunnan,China: Effects of Diet and Temperature

We studied seasonal variation in the activity budget of a habituated group of Nomascus concolor jingdongensis at Mt. Wuliang, Central Yunnan, China from March 2005 to April 2006 via scan sampling at 5-min intervals. The study site is near the northern extreme of the distribution of hylobatids, at high altitude with extreme seasonality of temperature and rainfall. During the day, feeding manifested a bimodal pattern of high activity levels in mid-morning and mid-afternoon, whereas resting reached a peak at midday, with proportionally less time used for traveling. Annually, the group spent an average of 40.0% of the time resting, 35.1% feeding, 19.9% traveling, 2.6% singing, 1.2% playing, and 1.3% in other activities. The proportion of time allocated to activities showed significant monthly variations and was influenced by the diet and temperature. Gibbons increased traveling and playing time and decreased feeding time when they ate more fruit, and they decreased traveling, singing, and playing time and increased feeding time when they ate more leaves. Moreover, when the temperature was low, the gibbons decreased time traveling and increased time resting. In summary, black-crested gibbons employed high-effort activities when they ate more fruit and energy-conservation patterns when they ate more leaves and in low temperature. Behavioral data from the site are particularly useful in understanding gibbon behavioral adaptations to different sets of ecological conditions.     

Effects of food and topography on ranging behavior of black crested gibbon (Nomascus concolor jingdongensis) in Wuliang Mountain, Yunnan, China

We studied the ranging behavior of a habituated group of black crested gibbons (Nomascus concolor jingdongensis) in a high, seasonal habitat on Mt. Wuliang, central Yunnan, China, between March 2005 and April 2006. Our results indicated that the total home range size for the study group was 129 ha, or 151 ha if the lacunae within the borders in which gibbons were not observed were included. This is a much bigger range size than that of other gibbon species. However, 69.7% of their activities occurred within 29 ha. The intensity of quadrant use was significantly correlated with the distribution of important food patches. The mean yearly daily path length was 1,391 m. Gibbons traveled farther when they spent more time feeding on fruit. To avoid often passing through ridges with little food, gibbons usually stayed in the same valley for successive days, and then moved on to another valley for another several days, which resulted in a concentrated ranging pattern.     

Gibbons under seasonal stress: the diet of the black crested gibbon (<Emphasis Type="Italic">Nomascus concolor</Emphasis>) on Mt. Wuliang,Central Yunnan,China

The diet of a habituated group of black crested gibbon (Nomascus concolor jingdongensis) was studied from March 2005 to April 2006 in the Wuliang Mountains, central Yunnan, China. Gibbons consumed 77 different plant species, one mammal-, two bird-, one lizard-, and two insect-species. Buds and leaves constituted 46.5% of the diet (21.0% vine leaves and buds, 19.2% tree leaves and buds, and 6.3% epiphyte leaves). Fruits, figs and flowers accounted for 25.5, 18.6 and 9.1% of the diet, respectively. There was marked seasonal variation in dietary proportions. Fruit varied from 0.3 to 82.7%; figs from 0 to 68.2%; tree leaves and buds from 1.5 to 83.3%; vine leaves and buds from 3.1 to 61.9%; and epiphyte leaves from 0 to 22.2% of the diet. Different food types dominated the diet in different months during the study period. The foli-frugivorous diet and extreme seasonal variation in the diet may be related to the harsh habitat of the study group.     

Sleeping Tree Selection of Cao Vit Gibbon (Nomascus nasutus) Living in Degraded Karst Forest in Bangliang,Jingxi, China

We studied the sleep‐related behavior of two Cao Vit gibbon (Nomascus nasutus) groups in Bangliang Nature Reserve in Jingxi County, China between January 2008 and December 2009 to test four hypotheses related to sleeping tree selection (predation avoidance, thermoregulation, food access, and range defense). Gibbons entered sleeping trees 88 ± SD 37 min before sunset before their main potential nocturnal predator become active. They usually moved rapidly and straight to sleeping trees and kept silent once settled. Over the course of the study, gibbon groups used many (87 and 57 per group) sleeping trees and reused them irregularly. They also tended to sleep in relatively tall trees without lianas, choosing small branches close to the treetop. These behaviors would make it difficult for potential terrestrial predators to detect and approach the gibbons. Therefore, these results strongly support the predation avoidance hypothesis. Gibbons tended to sleep closer to ridges than to valley bottoms and they did not sleep at lower elevations in colder months. They thus appeared not to select sleeping trees to minimize thermoregulatory stress. Gibbons very rarely slept in feeding trees, instead generally sleeping more than 100 m away from the last feeding trees of the day or the first feeding tree of the next morning. These patterns led us to reject the food access hypothesis. Lastly, we did not find evidence to support the range defense hypothesis because gibbons did not sleep in overlap areas with neighbors more often than expected based on the proportion of overlap and exclusively used areas. Am. J. Primatol. 74:998‐1005, 2012. © 2012 Wiley Periodicals, Inc.     

云南新平哀牢山西黑冠长臂猿分布与群体数量

西黑冠长臂猿(Nomascus concolor)主要分布于中国云南,滇中哀牢山是其重要分布区之一.哀牢山地处三州(市)六县(市),其中新平辖区是至今尚未全面开展西黑冠长臂猿种群数量与分布调查的地区.2009年11月—2010年1月,结合2007年和2009年对具有狩猎、放牧、林下资源采集经历的社区人员进行的访问调查,采用鸣声记数法对辖区内可能分布有长臂猿的区域进行了实地调查.调查结果显示,滇中哀牢山新平县辖区内分布有西黑冠长臂猿124群,其中85群栖息于国家级自然保护区及与其接壤的国有林内、30群栖息于县级自然保护区,9群栖息于自然保护区与国有林外的集体林中.可见,滇中哀牢山新平县辖区是迄今为止西黑冠长臂猿种群数量最大、分布最集中的区域,对西黑冠长臂猿保护具重要意义.但西黑冠长臂猿在该区域也呈现出斑块状集中分布的格局,以及种群分布密度和海拔梯度分布出现北高南低的趋势.     

Sleeping sites of black-and-white snub-nosed monkeys (Rhinopithecus bieti) at Baima Snow Mountain, China

Data on sleeping sites of a group of black-and-white snub-nosed monkeys Rhinopithecus bieti (Colobinae, Primates) were collected between April–July and September–December 2001 to try to determine the factors affecting site selection at Nanren (99°04'E, 28°34'N, Baima Snow Mountain, China). Sleeping trees were tall emergents (27.5±3.2 m) with large diameter at breast height (57.9±16.9 cm) and broad crown diameters (6.3±1.4 m), and significantly larger than other trees. The use of large sleeping trees could provide effective shelters from attacks of terrestrial and aerial predators, heavy snow and rain. Sleeping sites were changed nightly and located at significantly lower altitudes in winter months than in other seasons, which might be a thermoregulatory adaptation or related to snow covering. Such sites tended to be at the middle of sun-oriented slopes, possibly trading off predator avoidance and foraging efficiency.