Optimal group size and seasonal stress in ring-tailed lemurs (Lemur catta)

Adaptive explanations for social grouping assume that thereare fitness consequences associated with group size, and individualsmaintain membership in groups of favorable size to maximizefitness. Here I examine fecal cortisol concentrations as a hormonalmeasure of stress to assess the relative well-being of Lemurcatta in groups of different size and in seasons of normal andlow tamarind fruit availability. I test the hypotheses thatthere is an optimal group size at which cortisol is lowest andthat optimal group size changes in food-scarce conditions. Icollected 799 fecal samples from 87 individuals in seven free-rangingL. catta groups at Berenty Reserve, Madagascar, over a 1-yearperiod (August 1999–July 2000) and determined fecal cortisolconcentrations using a radioimmunoassay. Expressing these asresiduals from monthly population means to control for temporalfluctuations in cortisol concentration, I calculated mean fecalcortisol levels for each animal in seasons of normal and lowtamarind fruit abundance and over the entire year. Overall,females exhibited lowest mean cortisol levels in groups of intermediatesize, suggesting that there are benefits to maintaining membershipin these groups. Females in groups that were atypically largeor small for their habitat type had higher mean cortisol levelsthan typical groups. Cortisol levels increased in food-scarceconditions for larger groups, suggesting that intergroup competitiveadvantages do not outweigh intragroup feeding competition atthis time. Group size may be optimized for long-term averageconditions, and short-term stresses may intermittently alterthe costs associated with group size.     

The group size effect and synchronization of vigilance in the Tibetan wild ass

Vigilance behavior is considered as an effective strategy for prey species to detect predators.An individual benefits from living in a group by reducing the time spent being vigilant without affecting the probability of detecting a predator.However,the mechanism producing a decrease in vigilance with increasing group size is unclear.Many models of vigilance assume that group members scan independently of one another.Yet in recent studies,the other 2 patterns of vigilance,coordination and synchronization,were reported in some species.In 2 summers(2018 and 2019),we studied the group-size effect on vigilance and foraging of Tibetan wild ass in Chang Tang Nature Reserve of Tibet.We also tested whether individuals scan the environment independently,tend to coordinate their scans,or tend to synchronize their vigilance.The results showed that individuals decreased the time spent on vigilance with increasing group size,while increased the time spent foraging.Group members scanned the environment at the same time more frequently and there was a positive correlation between group members'behaviors,indicating that Tibetan wild asses tend to synchronize their vigilance.     

Number of neighbors instead of group size significantly affects individual vigilance levels in large animal aggregations

The group size effect states that animals living in groups gain anti‐predator benefits through reducing vigilance levels as group size increases. A basic assumption of group size effect is that all individuals are equally important for a focal individual, who may adjust its vigilance levels according to social information acquired from them. However, some studies have indicated that neighbors pose greater influences on an individual's vigilance decisions than other group members, especially in large aggregations. Vigilance has also been found to be directed to both predators (anti‐predation vigilance) and conspecifics (social vigilance). Central individuals might rely more on social vigilance than peripheral individuals. To test these hypotheses, we examined the effects of flock size, number of neighbors and position within a flock on vigilance and competition of greater white‐fronted goose Anser albifrons that form large foraging flocks in winter, controlling the effects of other variables (group identity, winter period and site). We found that individual vigilance levels were significantly affected by number of neighbors and position within a flock, whereas flock size showed no effect. Individuals devoted a large component of vigilance to nearby flock mates. Central individuals directed a relatively larger proportion of vigilance to monitor neighbors than peripheral ones, indicating that central individuals more relied on social information acquired from neighbors, possibly caused by the more blocked visual field of central individuals. Moreover, some social vigilance may function as conducting or preventing agonistic interactions since competition intensity was positively correlated with number of neighbors. Our study therefore demonstrate that the number of neighbors is more important than group size in determining individual vigilance in large animal groups. Further studies are still needed to unravel which neighbors pose greater influence on individual vigilance, and the factors that influence individuals to acquire information from their neighbors to adjust vigilance behaviors.     

Pen size and parity effects on maternal behaviour of Small-Tail Han sheep

The aim of this experiment was to study the effects of pen size and parity on maternal behaviour of twin-bearing Small-Tail Han ewes. A total of 24 ewes were allocated to a 2×2 design (six per pen), with parity (primiparous or multiparous) and pen size (large: 6.0×3.0 m; small: 6.0×1.5 m) as main effects at Linyi University, Shandong Province, China. Behaviour was observed from after parturition until weaning. All ewes were observed for 6 h every 5 days from 0700 to1000 h and from 1400 to 1700 h. Continuous focal animal sampling was used to quantify the duration of maternal behaviours: sucking, grooming and following as well as the frequency of udder accepting, udder refusing and low-pitched bleating. Oestradiol and cortisol concentrations in the faeces (collected in the morning every 5 days) were detected using EIA kits. All lambs were weighed 24 h after parturition and again at weaning at 35 days of age. The small pen size significantly reduced following (P<0.005), grooming (P<0.001) and suckling durations (P<0.05), as well as the frequency of udder refusals (P<0.001). However, there was a significant interaction with ewe parity, with decreased grooming and suckling in the small pen largely seen in the multiparous ewes (P<0.001). Independent of pen size, multiparous ewes accepted more sucking attempts by their lambs (P<0.05) and made more low-pitched bleats than primiparous ewes (P<0.001). Multiparous ewes had higher faecal oestradiol concentrations than primiparous ewes (P<0.001), and ewes in small pens had higher faecal cortisol levels compared with ewes in larger pens (P<0.001). As lambs increased in age, the duration of maternal grooming, following and suckling as well as frequency of udder acceptance and low-pitched bleating all declined, and the frequency of udder refusing increased (P<0.001 for all). Ewe parity, but not pen size, affected lamb weight gain during the period of observation (P<0.001). This is the first study to show that pen size, interacting with parity, can affect the expression of maternal behaviour in sheep during lactation. The study is also the first to report on the maternal behaviour of Chinese native sheep breeds (Small-Tail Han sheep), with implications for the production of sheep in China.     

Comparative physiological responses of stress in sheep selected for resistance or susceptibility to Haemonchus contortus in semi-arid tropical region during different seasons

The present study was conducted to assess the comparative physiological responses of stress in sheep selected for resistance or susceptibility to Haemonchus contortus in the semi-arid region during different seasons. A total of 20 adult Malpura sheep from resistant and susceptibility line were selected for study. The intensity of strongyle infection, blood biochemical and cortisol levels were estimated monthly basis while meteorological data were collected everyday throughout the year. The temperature-humidity index was the highest during dry-hot season. The faecal egg count increased significantly (p < 0.05) during June–August (rainfall season) in susceptible sheep as compared to resistant sheep. The plasma cortisol level increased significantly (p < 0.05) in both the groups during dry hot (March–May) season and it remained higher in the susceptible line till June–August. The total plasma protein reduced significantly during hot-humid with scantly rainfall (September–November) season as compared to other season in susceptible line. It is pertinent to conclude that though sheep remained in stress during dry hot (March–May) season, but effects of stress on the intensity of strongyle infection and blood biochemical were better tolerated by animals of resistant line during hot-humid season compared to susceptible line under semi-arid region.     

On how risk and group size interact to influence vigilance

Vigilance allows animals to monitor their surroundings for signs of danger associated with predators or rivals. As vigilance is costly, models predict that it should increase when the risk posed by predators or rivals increases. In addition, vigilance is expected to decrease in larger groups that provide more safety against predators. Risk and group size are thus two key determinants of vigilance. Together, they could have additive or interactive effects. If risk and group size interacted, the magnitude of the group‐size effect on vigilance would vary depending on the level of risk experienced, implying that the benefits of sociality in terms of vigilance vary with risk. Depending on the model, vigilance is predicted to decrease more rapidly with group size at low risk or at high risk. Little work has focused directly on the interaction between risk and group size, making it difficult to understand under which conditions particular interactive effects arise and whether interactive effects are common in natural systems. I review the vast literature on vigilance in birds and mammals to assess whether interactive effects between risk and group size are common, and if present, which pattern occurs more frequently. In studies involving predation risk, the greatest proportion reported no statistically significant interactive effects. In other cases, vigilance decreased with group size more rapidly at low or high risk in a similar proportion of studies. In studies involving risk posed by rivals (social risk), most documented a more rapid decrease in vigilance with group size at low than at high risk, as predicted if the need to monitor rivals increases in larger groups. Low statistical power to detect interactive effects might have been an issue in several studies. The absence of interactive effects, on the other hand, might suggest constraints or limits on the ability of animals to adjust vigilance to current risk or group sizes. Interactive effects on vigilance have implications for the evolution of sociality and for our understanding of the phenotypic plasticity of predator‐ and competitor‐induced defences and deserve more attention in future studies.     

Vigilance and group size in impala (Aepyceros melampus Lichtenstein): a study in Nairobi National Park, Kenya

One of the advantages of living in groups is that individuals may need to be less vigilant, allowing them more time for other important activities, such as foraging. This relationship between group size and per cent time spent being vigilant was investigated by observing impala in Nairobi National Park, Kenya. Three types of individual were observed: females, territorial males and bachelor males. Only females showed the predicted negative relationship between per cent vigilance and herd size. Both types of male showed no significant change of vigilance with increasing group size. There was no difference in levels of vigilance in open or closed habitats and no difference in vigilance between herds ‘alone’ and herds with other species that might have provided ‘extra eyes’.     

Group size effect on vigilance: evidence from Tibetan gazelle in Upper Buha River, Qinghai-Tibet Plateau

Tibetan gazelle Procapra picticaudata is a threatened and endemic species to the Qinghai-Tibet Plateau. With the method of group scan sampling, we observed the behaviours of males and females of the gazelle in the two summers of 2005 and 2006, in order to test the group size effect on group vigilance. We found that male gazelles were significantly more vigilant than the females at both group scan level (percentage of individuals scanning during a session) and group scan frequency (percentage of intervals with at least one individual scanning). We also found a negative correlation between group scan level and group size and a positive correlation between group scan frequency and group size, showing the group size effect on vigilance was testified in Tibetan gazelle. The predation factor might be the main driving force for the group size effect.     

Group size affects the metabolic rate of a tropical beetle

ABSTRACT. . Dormancy is metabolically very costly for insects and other poikilotherms of the tropics because energy reserves are rapidly utilized at high temperature. Yet, the beetle Stenotarsus rotundus Arrow spends up to 10 months each year in a tropical lowland forest as a non-feeding adult, dependent upon its fat reserve as an energy source. While in this dormant state, the beetles aggregate in huge numbers and the size of the group greatly influences the metabolic rate. As group size increases, the metabolic rate decreases, and the minimum rate observed in the field (22 μ l O₂ g^-1 h^-1) is estimated to occur when group size reaches about 300 individuals. A decrease in metabolic rate can also be achieved by elevating relative humidity, thus suggesting that one function of the aggregation is to increase humidity within the group and thereby decrease metabolic rate.     

Group formation of female pigtail macaques (Macaca nemestrina)

Human epidemiological studies have suggested that social variables can modulate the effects of stress on the immune system, and this concept has been gaining increasing attention with positive results emerging from empirical studies using nonhuman primates over the last two decades. Results from a previous study in rhesus monkeys suggested that receiving grooming positively affected recovery of T-helper and T-suppressor cells following the initial stress associated with group formation, and this co-varied with high dominance rank. Thus, the present study was undertaken in order to determine: (1) if the stress effect of formation could be replicated in another species and (2) if social behaviors or dominance rank, given that formation is a stressor, might independently correlate with physiological recovery from the stressor. Eight adult female pigtail macaques were moved from individual cages and simultaneously introduced into an outdoor enclosure along with an adult male, while eight weight-matched controls remained in individual caging. Behavioral data were collected during the introduction and over 4 weeks thereafter. Blood samples were collected prior to and at intervals for 4 weeks following formation. Compared to control subjects, the test subjects showed an increase in basal cortisol secretion (+28.9%) and a significant decrease in T-helper cells (-33.6%), T-suppressor cells (-30.8%), and B cells (-22.5%), while there was a significant increase in white blood cells (+29.5%) 24 hr following formation. When dominance rank and seven behavioral categories were analyzed, only the frequency of receiving grooming significantly predicted change, with animals who received a greater frequency of grooms showing a lesser negative percent change from baseline in the absolute number of T-helper cells 1 week following formation. The establishment of a dominance hierarchy, apparent within 1 week, was accomplished with no serious fighting and a complete absence of wounding or trauma, suggesting that psychosocial stress was responsible for the physiological changes observed. © 1996 Wiley-Liss, Inc.     

Wool cortisol is a better indicator of stress than blood cortisol in ewes exposed to heat stress and water restriction

This study investigated the effect of water restriction on wool and blood cortisol concentrations and water consumption patterns in heat-stressed sheep. Nine Corriedale female sheep (average BW=43±6.5 kg) were individually fed diets based on maintenance requirement in metabolic crates. They were assigned to three treatments according to a Latin square design (3×3) for three periods with a 21-day duration for each period (nine sheep per treatment). Treatments included free access to water (FAW), 2 h water restriction (2hWR) and 3 h water restriction (3hWR) after feeding. Average temperature–humidity index in the experimental room was 27.9 throughout the experiment that defines heat stress conditions. Wool samples were taken at the end of each period on day 21. No differences were found in cortisol concentration in each fragment (dried, washed and residual extract) of wool (P<0.05). Total wool cortisol concentration was higher in the 3hWR group than the other treatments (P<0.05). Blood cortisol was not different among the treatments (P>0.05) and resulted in higher variable data compared with wool cortisol. Blood neutrophils and neutrophil/lymphocyte ratio suppressed in FAW and 3hWR groups compared with the 2hWR group (P<0.05). The duration of water consumption recorded after feeding in the 3hWR group was higher than in the 2hWR group when recorded in the afternoon (P<0.01). Water consumption rate was higher in the 3hWR group than in the 2hWR group (P<0.01). However, total water consumed was lower in the 3hWR group compared with other treatments (P>0.05). It can be concluded that wool cortisol provides more precise and accurate data than blood cortisol during heat stress conditions. Water restriction for 3 h after feeding can act as a stressor and is critical for sheep during heat stress as the consumption of water decreases with restriction.     

The effect of shearing procedures on blood levels of growth hormone, cortisol and other stress haematochemical parameters in Sarda sheep

The aim of this research was to investigate how growth hormone (GH) cortisol and some haematochemical parameters could be modified by the stress caused by the stages of shearing in Sarda breed sheep. Five groups of 10 sheep each were formed. Group A, only separated from the flock; Group B, only tied; Group C, both tied and shorn (animals in these three groups were ewe lambs shorn for the first time); Group D, adult females both tied and shorn; and Group E, adult entire males both tied and shorn (animals in these two groups had been shorn previously). Five blood samples were taken from each animal: the day before treatment (first sample); at the start of the treatment (second sample); in the middle of shearing for Groups C, D and E, 10 min after separation in Group A and 10 min after tying in Group B (third sample); at the end of treatment (fourth sample); and on the day after treatment (fifth sample). Plasma GH levels showed a decrease (P < 0.01) in Groups A, B, C and D during treatment (third and fourth samples), while Group E only at the end of shearing (fourth sample). In the third sample, the highest GH levels were recorded for Group E (P < 0.05), while it was recorded in the fourth sample for Groups A and E (P < 0.05). Cortisol levels showed a clear increase (P < 0.01) in all groups during treatment, but Group A showed a decrease in the fourth sample in comparison to the third sample. Males in the second, third and fourth sample and Group A only in the fourth sample showed lower cortisol levels when compared with the other groups (P < 0.05). Plasma glucose levels showed an increase (P < 0.01) in all groups during treatment but Groups B, C and E showed the highest values (P < 0.05). Magnesium (Mg) showed an increase in all groups in the third and fourth sample, while sodium (Na), in the same samples, only in Groups B, C and D. Potassium (K) values showed a significant decrease (P < 0.05) only in Groups C and D at the end of shearing. These results show that GH secretion is influenced by all the stress procedure: separation, tying and shearing. Shearing, even if necessary for animals, causes a significant change of the blood parameters involved in the stress response.     

The relationship between ecological segregation and sexual body size dimorphism in large herbivores

Ecological segregation (sexual differences in diet or habitat use) in large herbivores has been intimately linked to sexual body size dimorphism, and may affect both performance and survival of the sexes. However, no one has tested comparatively whether segregation occurs at a higher frequency among more dimorphic species. To test this comparatively, data on sex-specific diet, habitat use and body size of 40 species of large herbivores were extracted from the literature. The frequency of ecological segregation was higher among more dimorphic herbivores; however, this was only significant for browsers. This provides the first evidence that segregation is more common among more dimorphic species. The comparative evidence supported the nutritional-needs hypothesis over the incisor breadth hypothesis, as there was no difference in frequency of segregation between seasons with high and low resource levels, and since segregation was also evident among browsers. Whether the absence of a correlation between ecological segregation and level of sexual body size dimorphism for intermediate feeders and grazers is due to biological differences relative to browsers or to the fact that the monomorphic species included in the analysis were all browsers is discussed. Received: 18 August 1999 / Accepted: 31 January 2000     

Group size variability in primates

The effects of mean troop size, diet, territoriality, and habitat upon temporal variability of group size in primates were investigated using variance functions relating mean group size and temporal variability. Two different types of variability were described: (1) within group variability where a single troop was followed over a given period of time; and (2) between group variability where the author(s) did not distinguish one troop from the other. In the second category, CV (SD/mean) as an index of temporal group size variability proved to be dependent on mean group size among the Cercopithecidae. Large groups are more unstable in size than small ones. In the Cebidae, variability was independent of mean group size and therefore large groups are as variable through time as small ones. Ecological factors showed no effects on the observed level of between group variability. Within group variability was found to be smaller than the level of between group variability in all species tested. The results are related to social organization and to the degree of feeding interference observed within and between troops. Future practical applications for our results are considered.     

Group size,communication, and familiarity effects in foraging human teams

Exploitation of food resources that are dispersed in time and space has been crucial to the evolutionary success of humans. Recent experimental work has shown that an absence of communication impairs decision‐making in a foraging task. Here, we found that individuals in larger teams were more likely to reach group consensus and were more accurate and efficient foragers. Individuals in larger teams were also more likely to gesture to one another, while levels of verbal exchange were not significantly different in small and large groups. At last, teams in which individuals reported that they knew one another and rated team members as helpful and information‐seeking were more accurate in their foraging. Overall, our findings offer experimental evidence that larger, communicating, familiar teams are quicker and more accurate foragers. We therefore suggest that complex communication within socially bonded relationships may have been important to the ecological success of the human lineage.     

Behavioral response of 12 ungulate species in captivity to the presence of humans

Twelve species of captive ungulates were studied to determine behavioral responses to the presence of a zookeeper within the exhibit and in front of the exhibit, with and without zoo visitors present. Significant differences in behavior occurred between species for nearly all behaviors observed. A significantly greater occurrence of vigilance and approach behavior was directed toward the zookeeper while within the exhibit relative to when the zookeeper stood in front of the exhibit. A significantly lower occurrence of eating or drinking occurred when the zookeeper was inside the exhibit. Significant differences occurred across size categories in the occurrence of approaching the zookeeper. The most frequently scored behavior was visual orientation. Statistically significant differences in the occurrence of visual orientation directed by females toward the zookeeper existed across size category, with more vigilance by species with larger body size. A statistically higher occurrence of vigilance toward the zookeeper was directed by female ungulates but not by males when the zoo was closed to the public. Although no statistical significance was found regarding the intraspecific vigilance of males, data on females revealed significant differences across species and across size categories. When data regarding vigilance toward the public were analyzed, statistically significant differences existed between species for females only. Likewise, when data regarding interspecific vigilance were examined, statistically significant differences were found across size categories for females, but not for males. The potential roles of vigilance in the wild are discussed in reference to its role in captivity.     

Conspecific Influences on Vigilance Behavior in Wild Chimpanzees

Diurnal primates rely on visual monitoring behavior to collect various kinds of ecological and social information. Vigilance behavior is monitoring specifically to detect external threats. Previous studies of vigilance behavior were focused mainly on the influence of predation threats, whereas the influences of conspecific factors, such as intragroup threats, have been relatively unstudied. Individual vigilance is predicted to be inversely related to the group size or the number of individuals nearby if the main target of the vigilance is a predation threat and positively related if the main target of the vigilance is a conspecific threat. I studied wild chimpanzees (Pan troglodytes schweinfurthii) in Mahale Mountains National Park, Tanzania, and measured the vigilance duration when they are resting on the ground via 2-min focal observation. In both males and females, vigilance duration increased as the number of individuals nearby increased. This result agrees with the idea that the chimpanzees are vigilant toward other group members. In addition, maternal vigilance monitors and protects the safety of dependent offspring as the duration of maternal vigilance was longer when a dependent infant was separated from its mother than when the offspring was in contact with its mother. The results indicate that the vigilance behavior in wild chimpanzees was affected by conspecific factors.