Moving calls: a vocal mechanism underlying quorum decisions in cohesive groups

Members of social groups need to coordinate their behaviour when choosing between alternative activities. Consensus decisions enable group members to maintain group cohesion and one way to reach consensus is to rely on quorums. A quorum response is where the probability of an activity change sharply increases with the number of individuals supporting the new activity. Here, we investigated how meerkats (Suricata suricatta) use vocalizations in the context of movement decisions. Moving calls emitted by meerkats increased the speed of the group, with a sharp increase in the probability of changing foraging patch when the number of group members joining the chorus increased from two up to three. These calls had no apparent effect on the group's movement direction. When dominant individuals were involved in the chorus, the group's reaction was not stronger than when only subordinates called. Groups only increased speed in response to playbacks of moving calls from one individual when other group members emitted moving calls as well. The voting mechanism linked to a quorum probably allows meerkat groups to change foraging patches cohesively with increased speed. Such vocal coordination may reflect an aggregation rule linking individual assessment of foraging patch quality to group travel route.     

Who cares who calls? Selective responses to the lost calls of socially dominant group members in the white‐faced capuchin (Cebus Capucinus)

In many social mammals and birds, soft vocalizations are habitually produced during dispersed moving and foraging, the function being to maintain contact and regulate spacing between group members. In some species, much louder calls are given sporadically by specific individuals when they become separated from the group, or 'lost'. The function of these calls has seldom been specifically tested, particularly among social primates, but is often assumed to involve regaining contact with the group based on a combination of individually distinctive calls and antiphonal responses to them from within the group. To test these assumptions, we conducted research on two groups of white-faced capuchins (Cebus capucinus) in Costa Rica. We analyzed 82 bouts of 'lost' calls given by 13 different adult individuals when separated from the group and the antiphonal responses they elicited. Lost calls were individually distinctive and were answered in 35% of calling episodes. Answers were selective: dominant males and females were answered more than were subordinate callers of either sex. As a result, dominant callers relocated and returned to the group more quickly than did subordinate callers. We discuss the potential proximate motivations for, and ultimate benefits of, such selective answering of dominant group members.     

Individuals in foraging groups may use vocal cues when assessing their need for anti-predator vigilance

Many studies of social species have reported variation in the anti-predator vigilance behaviour of foraging individuals depending on the presence and relative position of other group members. However, little attention has focused on how foragers assess these variables. It is commonly assumed that they do so visually, but many social species produce frequent calls while foraging, and these 'close' calls might provide valuable spatial information. Here, we show that foraging pied babblers (Turdoides bicolor) are less vigilant when in larger groups, in the centre of a group and in closer proximity to another group member. We then show that foragers are less vigilant during playbacks of close calling by more individuals and individuals on either side of them when compared with calls of fewer individuals and calls on one side of them. These results suggest that foragers can use vocal cues to gain information on group size and their spatial position within a group. Future studies of anti-predator vigilance should consider the relative importance of both visual and vocal monitoring of group members.     

Effects of caller activity and habitat visibility on contact call rate of wild Japanese macaques (Macaca fuscata)

A major function of contact calls in nonhuman primates is to maintain spatial cohesion among individuals in a group. The risks of spatial/visual separation from the group are likely to affect auditory contact behavior, in particular by increasing the call rate. We tested whether the risk of separation influences coo call emission by investigating the variation in call rate among behavioral contexts in two wild populations of Japanese macaques (Macaca fuscata). We focused on caller activity and the degree of visibility within the habitat as primary potential factors mediating call rate. We first estimated the habitat visibility of the two research sites at Yakushima Island (YK) and Kinkazan Island (KZ), Japan. The habitat visibility of YK was significantly more restricted than that of KZ. We then compared the call rate of 20 adult and 12 juvenile female macaques between the two wild populations to examine the potential effects of environmental differences. Both populations had a lower call rate during grooming than during feeding and moving, which are behaviors associated to higher interindividual distances. The call rate of YK adult females was significantly greater than that of both juveniles and KZ adult females, independently of activity. The call rate increased as macaques matured in the YK population, but not in the KZ population, suggesting that different developmental processes involved in contact calling of the two populations. Our findings suggest that separation risk influences call rate, and also imply a possibility of social influence that social structure change effects on the call rates.     

Maintaining social cohesion is a more important determinant of patch residence time than maximizing food intake rate in a group-living primate,Japanese macaque (Macaca fuscata)

Animals have been assumed to employ an optimal foraging strategy (e.g., rate-maximizing strategy). In patchy food environments, intake rate within patches is positively correlated with patch quality, and declines as patches are depleted through consumption. This causes patch-leaving and determines patch residence time. In group-foraging situations, patch residence times are also affected by patch sharing. Optimal patch models for groups predict that patch residence times decrease as the number of co-feeding animals increases because of accelerated patch depletion. However, group members often depart patches without patch depletion, and their patch residence time deviates from patch models. It has been pointed out that patch residence time is also influenced by maintaining social proximity with others among group-living animals. In this study, the effects of maintaining social cohesion and that of rate-maximizing strategy on patch residence time were examined in Japanese macaques (Macaca fuscata). I hypothesized that foragers give up patches to remain in the proximity of their troop members. On the other hand, foragers may stay for a relatively long period when they do not have to abandon patches to follow the troop. In this study, intake rate and foraging effort (i.e., movement) did not change during patch residency. Macaques maintained their intake rate with only a little foraging effort. Therefore, the patches were assumed to be undepleted during patch residency. Further, patch residence time was affected by patch-leaving to maintain social proximity, but not by the intake rate. Macaques tended to stay in patches for short periods when they needed to give up patches for social proximity, and remained for long periods when they did not need to leave to keep social proximity. Patch-leaving and patch residence time that prioritize the maintenance of social cohesion may be a behavioral pattern in group-living primates.     

The time budget of a group of wild woolly monkeys (Lagothrix lagotricha)

I studied a group of woolly monkeys (Lagothrix lagotricha)for 1800 hr from early 1985 until September 1987 near the lower Apaporis river of the Vaupés Department in the eastern Colombian Amazon. The group had an activity budget of 29.9% rest, 38.8% move, 25.8% forage, and 5.5% other activities during 720 hr of observations evenly spaced throughout this year. The only significant change in these activities during the study period was social activity, which increased as a function of increased food supply. Average diurnal activities of resting and moving are negatively correlated, and each activity show significant changes during the day, while foraging show no significant change. The activity budgets of the adult males and females are not significantly different, but the activities of the juveniles and infants showed much more social activity than the adults. High moving scores may be correlated with the comparative infertility of the soils of the Caparú forest, which require increased effort to maintain the energy levels necessary for maintenance and reproduction in Lagothrix.     

The time budget of a group of wild woolly monkeys (Lagothrix lagotricha)

I studied a group of woolly monkeys (Lagothrix lagotricha)for 1800 hr from early 1985 until September 1987 near the lower Apaporis river of the Vaupés Department in the eastern Colombian Amazon. The group had an activity budget of 29.9% rest, 38.8% move, 25.8% forage, and 5.5% other activities during 720 hr of observations evenly spaced throughout this year. The only significant change in these activities during the study period was social activity, which increased as a function of increased food supply. Average diurnal activities of resting and moving are negatively correlated, and each activity show significant changes during the day, while foraging show no significant change. The activity budgets of the adult males and females are not significantly different, but the activities of the juveniles and infants showed much more social activity than the adults. High moving scores may be correlated with the comparative infertility of the soils of the Caparú forest, which require increased effort to maintain the energy levels necessary for maintenance and reproduction in Lagothrix.     

不同天气条件下青头潜鸭越冬期行为的响应模式

2018年11—12月在河南民权国家湿地公园,采用定点观察法、扫描取样法对越冬青头潜鸭在不同天气条件下的行为响应模式进行研究。结果表明: 在晴天,静息、飞行、运动是青头潜鸭越冬期的优势行为(65.5%),觅食、修整是次要行为(31.9%);行为节律表现为上午觅食、中午静息、下午觅食修整;飞行通常发生在觅食高峰前,运动行为多与其他行为相伴,其时间分配与觅食呈正响应,与静息呈负响应。相较于晴天,阴雨天青头潜鸭增加了对静息和觅食的时间分配,静息、觅食和运动为阴雨天的主要行为(76.5%),修整、飞行为次要行为(20.3%);觅食和静息的高峰时段后移,傍晚时达到高峰,飞行和修整时间显著减少。与晴天相比,雾霭天青头潜鸭静息、觅食、运动和修整的时间分配增加,飞行行为减少;觅食高峰后移至中午和下午,静息高峰后移至下午;静息、运动和觅食为雾霭天的优势行为(70.6%),修整和飞行为次要行为(27.5%)。研究表明,不同天气条件下,民权湿地公园青头潜鸭的越冬期行为在时间分配、行为节律、优势行为方面存在一定的响应;阴雨天和雾霭天青头潜鸭会分配更多的静息和觅食时间,并通过增加能量摄入和减少耗能行为应对恶劣天气。     

A preliminary study of vocal communication in wild long-tailed macaques (Macaca fascicularis). II. Potential of calls to regulate intragroup spacing

This study is a preliminary assessment of the potential of long-tailed macaque (Macaca fascicularis) calls to operate in systems of within-group spacing. Covariation in the rate of occurrence of calls with party spread, size, and activity among wild individuals of one group suggested that four classes of calls may function in intragroup spacing. Two of them are “clear” calls of long duration and pronounced frequency modulation. Calling rate increased with party spread for low- and high-frequency variants of these calls during resting and feeding respectively, suggesting possible utility in maintenance of spatial relations over moderately long distances. A third “harsh” call was negatively correlated with party spread during foraging and may thus function to increase dispersion among foraging individuals. Another harsh call with a tonal onset was unique among all calls in the vocal repertoire in being more frequently performed by lone, isolated individuals than by macaques accompanied by others, suggesting a possible function in reestablishing contact that has been severed. The functional significance of these calls with respect to their acoustic structures is discussed. Macaques that use calls to regulate intragroup spacing can control communication distance and direction by their choice of acoustically different vocalizations. This choice may be affected not only by varying environmental constraints on sound transmission, but also by social and ecological factors such as intragroup competition.     

在不同状态和生境复杂度中大蹄蝠回声定位叫声的可塑性

蝙蝠回声定位声波的可塑性对其适应不同状态、生境以及捕食和社会交流具有重要的作用。为进一步研究大蹄蝠的回声定位声波在不同状态和生境下的可塑性,通过室内行为实验,对大蹄蝠在4 种不同状态(室内飞行、静息、布袋内和手持)和4 种不同生境复杂度(室外、室内0 棵树、室内1 棵树、室内5 棵树)条件下飞行的回声定位声波特征进行研究。结果表明:大蹄蝠的回声定位声波为CF - FM 型,通常连续发出2 - 4 个脉冲组成一个脉冲组。对比大蹄蝠在4 种不同状态下的回声定位叫声发现,主频按静息、布袋内、手持、飞行的顺序依次降低,后端FM 频宽则按手持、布袋内、飞行和静息的顺序依次变短;而脉冲间隔和脉冲时程则均按静 息、飞行、布袋内、手持的顺序依次增加。对比大蹄蝠在4 种不同生境复杂度中飞行的回声定位叫声发现,主频按室外、室内0 棵树、室内1 棵树、室内5 棵树依次提高,而脉冲时程及脉冲间隔则依次缩短;室外放飞条件下的后端FM 频宽比室内飞行的短。研究结果说明,大蹄蝠在不同状态、不同生境复杂度条件下的回声定位叫声具有明显的可塑性和生境适应性。     

Experimental evidence for group hunting via eavesdropping in echolocating bats

Group foraging has been suggested as an important factor for the evolution of sociality. However, visual cues are predominantly used to gain information about group members'' foraging success in diurnally foraging animals such as birds, where group foraging has been studied most intensively. By contrast, nocturnal animals, such as bats, would have to rely on other cues or signals to coordinate foraging. We investigated the role of echolocation calls as inadvertently produced cues for social foraging in the insectivorous bat Noctilio albiventris. Females of this species live in small groups, forage over water bodies for swarming insects and have an extremely short daily activity period. We predicted and confirmed that (i) free-ranging bats are attracted by playbacks of echolocation calls produced during prey capture, and that (ii) bats of the same social unit forage together to benefit from passive information transfer via the change in group members'' echolocation calls upon finding prey. Network analysis of high-resolution automated radio telemetry confirmed that group members flew within the predicted maximum hearing distance 94±6 per cent of the time. Thus, echolocation calls also serve as intraspecific communication cues. Sociality appears to allow for more effective group foraging strategies via eavesdropping on acoustical cues of group members in nocturnal mammals.     

The Huh Vocalization of White-faced Capuchins: a Spacing Call Disguised as a Food Call?

White-faced capuchins, (thus capuchins, predictably emit huh vocalizations at high rates within dense fruit patches. We sought to determine why white-faced capuchins at the La Selva Biological Station, Costa Rica produce these food-associated calls. Here we analyze the contexts in which this intra-group vocalization was emitted, including the spatial responses elicited from other troop members. A cumulative 26.6 h of continuous focal samples and 3314 spectrograms (including 1643 huhs) were analyzed from a study troop with 16 focal subjects. The mean individual rate of huhs was greater (1) during foraging versus nonforaging activities; (2) during fruit foraging compared to both visual searching for foraging sites and foraging for arthropod prey; and (3) when the nearest neighbor was within a 10 m radius of the focal animal compared to when the nearest neighbor was at greater distances. A huh also predicted a significant increase in nearest-neighbour distance; on average, mean nearest-neighbor distance increased 3 m within 2 min following a huh vocalization. Null models of change in mean nearest-neighbor distance over time were generated from the original data set by treating predetermined time points (140 s intervals) in the focal recordings as if those points marked instances at which huhs were produced by the focal subject. No significant alterations in nearest-neighbor distance were detected within time lags up to 100 s in these null models, supporting the conclusion that huhs are causally linked with subsequent increases in nearest-neighbor distances. Huhs were most evident when capuchins were within dense fruit patches, but these calls were produced across all foraging contexts. Our results suggest that huhs may not be food calls in the usual sense (i.e. informing others of the location of food sources to be shared), but may be more appropriately described as spacing calls. Huhs probably act to increase foraging efficiency by reducing overlap in foraging areas with other troop members.     

Associations between adult spectral tarsiers

While anecdotal observations of gregarious behavior in nocturnal prosimian primates are common, most anthropologists continue to refer to them as solitary, perhaps based on the assumption that the occasional social interactions observed via ad libitum methods represent random chance encounters and not patterned social interactions. In this paper, I test the null hypothesis that nocturnal encounters between spectral tarsier (Tarsius spectrum) group members, outside of the sleeping tree, are the result of chance. Three male‐female pairs were radio‐collared and observed over a 4‐month period, using continuous focal animal sampling at the Tangkoko Nature Reserve (Sulawesi, Indonesia). Using Waser's random gas model, I found that spectral tarsiers spent more time in proximity to other group members than expected by chance, given the size of their home range and nightly path length. Adult group members spent 11% of the night in physical contact and an additional 17% of the night within a 10‐m radius of one another. Spectral tarsiers were also observed to significantly increase the amount of time spent foraging when located less than 10 m from another group member. Individuals foraging in proximity to another adult group member had lower insect capture rates compared to individuals who were not foraging in proximity to another adult group member. If living in a group is costly to these tarsiers' foraging efficiency, then why don't they actively avoid one another when foraging? One situation in which it might benefit tarsiers to be gregarious is high predation pressure. Preliminary results suggest that predation pressure by snakes may be the most likely factor selecting for the tarsiers to forage in proximity. Am J Phys Anthropol 128:74‐83, 2005. © 2005 Wiley‐Liss, Inc.     

Effects of contextual and social variables on contact call production in free-ranging ringtailed lemurs (Lemur catta)

Ringtailed lemurs (Lemur catta)often emit a characteristic vocalization when resting as a troop. This “meow call” is believed to serve an intragroup contact function. I investigated the effects of behavioral context and proximity on call production and acoustic features of the vocalization in a free-ranging troop in the Berenty Reserve, Madagascar. When group members were dispersed, the meow call was given frequently, and the acoustic features of the call probably increased the locatability of the emitter. I evaluated social relationships among the troop members from grooming interactions and spacing data. I also studied the call exchange network among the troop members, the results of which indicate that calls are exchanged between the two subgroups of females, which they connect.     

Social influences on foraging behaviour of the Red-cheeked Cordon Bleu Uraeginthus bengalus Estrildidae

Red-cheeked Cordon Bleus U. bengalus forage on bare patches of ground for fallen grass seeds and termites in particular home ranges. Foraging is confined to shaded areas in the middle parts of the day but birds exploit places exposed to the full sun in the early mornings and late afternoons. They mostly occur in pairs but often aggregate to form temporary foraging groups. Although birds in groups spend less time vigilant than those foraging alone, they react sooner than them to approaching danger. Birds foraging in pairs peck more rapidly than those in other situations and it is suggested that one advantage of forming permanent pair bonds is that paired birds forage more efficiently than unpaired ones in preparation for the coming breeding season. Flight calls, contact calls and alarm calls are given in specific behavioural contexts. Birds make contact calls when they become separated from their mates and evidently use them to re-locate one another. It is shown experimentally that members of pairs can recognize one another's contact calls.     

Flock density, social foraging, and scanning: an experiment with starlings

Social foraging differs from individual foraging because italters both resource availability and the forager's behavior.We examined responses of starlings to the presence of conspecificsby manipulating foraging-group density experimentally, whileensuring that each subject's foraging opportunities were unchanged.To do this, we used individuals foraging simultaneously in fourbottomless enclosures placed at various separations in naturalforaging grounds. We measured foraging and scanning intensityand qualitative aspects of scanning of focal individuals. Additionally,we examined the temporal distribution of scanning between individuals.The focal individual analysis showed that (1) food-searchingactivity increased, while time spent scanning, time off theground and scanning bout length decreased with flock density;(2) food finding per unit of searching effort increased withdensity; (3) head orientation during scanning was sensitiveto companions' proximity: heads pointed away from the companionsat close distance, toward them at intermediate distance, andwas random farther away. The analysis of the (temporal overlappingin scanning) temporal distribution of scanning for the groupshowed that scanning was significantly synchronized when companionswere adjacent to each other but was not significantly differentfrom random at further separations. We conclude that behavioralresponses of individuals to the presence of others generateimportant changes in foraging performance even in the absenceof physical interference and, more generally, that assessingthe mechanisms that control the behavior of group members atdifferent flock densities offers a way to understand the functionaland ecological significance of foraging aggregations.     

Vocal Coordination of Group Movement by Green Woodhoopoes (Phoeniculus purpureus)

Communication signals are used by many species to maintain group cohesion when moving over larger areas. Groups of green woodhoopoes (Phoeniculus purpureus) generally move around their territory as a close‐knit unit. Dominant individuals were more likely than subordinates to initiate movement to a new foraging site, but there was no intersexual difference. Dominants were also more likely than subordinates to be followed immediately. Vocalizations were shown to play an important role in mobilization: in the thick forests inhabited by woodhoopoes, visual cues to coordinate movement are likely to be less successful. When responding to the rallying call of a neighboring group, dominants and subordinates were equally likely to lead, as were males and females. As other group members followed immediately on most of these occasions, vocalizations were less important in this context than when moving to a new foraging site.     

Compete to Play: Trade-Off with Social Contact in Long-Tailed Macaques (Macaca fascicularis)

Many animal species engage in various forms of solitary object play, but this activity seems to be of particular importance in primates. If playing objects constitute a valuable resource, and access to such objects is limited, a competitive context may arise. We inserted a unique toy within a mini-colony of long-tailed macaque (Macaca fascicularis) and compared their behaviors to sessions without playing object. An automatic color-based 3D video device was used to track the positions of each animal and the toy, and this data was categorized into 5 exclusive behaviors (resting, locomotion, foraging, social contact and object play). As expected, the delay to first access to the object reflected the hierarchy of the colony, indicating that a competition took place to own this unique resource of entertainment. In addition, we found that the amount of object play was not correlated with social or foraging behavior, suggesting independent motivational mechanisms. Conversely, object playing time was negatively correlated with idling time, thus indicating its relation to pastime activities. Interestingly, the amount of social contacts in the group was significantly reduced by the heightened competitive context, suggesting that competitors are more likely to be perceived as potential threat requiring caution, as shown in humans. Experimental manipulation of competitive contexts in primates reveals common mental processes involved in social judgment, and shows that access to valuable resources can be a sufficient cause for variations in group cohesion.     

Response of foraging group members to sentinel calls in suricates,Suricata suricatta

In the suricate (Suricata suricatta), a cooperatively breeding mongoose, one individual typically watches out for predators while the rest of the group is foraging. Most of the time these sentinels announce their guarding duty with special vocalizations. The response of foraging group members to these calls was investigated by analysing observational data, and by performing playback experiments. The use of special calls by sentinels, and the responses of the foraging group members to them, suggest that the coordination of vigilance is strongly influenced by vocal communication. Sentinel calls decreased the time spent alert by the foraging group members. Other group members were less likely to go on guard when a sentinel was vocalizing. Both the proportion of time during which guards overlapped, and the proportion of time the group was unprotected without a guard, decreased when sentinels announced their duty, due to better coordination of the rotation of sentinels. Suricates, however, do not appear to use sentinel calls to mediate a strict rotation of guarding duty.     

白眼潜鸭秋季行为时间分配及活动节律

2012年7~11月,采用焦点动物取样法和瞬时扫描法,在乌鲁木齐近郊水域白湖,对白眼潜鸭（Aythya nyroca）秋季行为进行实地观察研究。制定出白眼潜鸭行为谱,各种行为有取食、保养、运动、休息、警戒、其他,共6类14种。昼间各行为时间分配依次为取食（56.7%）、运动（13.8%）、保养（11.8%）、休息（11.2%）、警戒（5.1%）、其他（1.4%）。其中取食、休息、运动行为有明显的节律性变化。休息行为以家庭为单位聚集在一起进行,取食行为则分开进行。7~9月份7：00~8：00时、18：00~19：00时休息行为出现峰值,10~11月份8：00~9：00时、17：00~18：00时出现峰值;7~9月份取食行为在8：00~9：00时、17：00~18：00时有明显的峰值,10~11月份在10：00~11：00时和15：00~16：00时出现峰值;7~11月份运动行为均在中午达到峰值。白眼潜鸭7~9月份与10~11月份觅食行为、休息行为和保养行为差异极显著（P<0.01）,运动行为无显著性差异（P >0.05）。迁徙前（10月中旬）,白眼潜鸭有集群现象,数量最多达到37只,10月下旬种群集体迁徙。秋季白眼潜鸭主要时间用于取食,意图为了迁徙或越冬积累能量,集群迁徙有利于种群防御天敌。