Predation risk, gender and the group size effect: does elk vigilance depend upon the behaviour of conspecifics?

Many animals benefit from the presence of conspecifics by reducing their rate of scanning for predators while increasing their time spent foraging. This group size effect could arise from a decreased perception of individual risk (dilution hypothesis) and/or an increased ability to detect predators (detection hypothesis). We compared individual and group vigilance of Rocky Mountain elk, Cervus elaphus, in three regions of Yellowstone National Park, Wyoming, U.S.A. that varied in their encounter frequency with coyote, Canis latrans, grizzly bear, Ursus arctos, and grey wolf,Canis lupus , predators. Adult females without calves increased scanning and decreased foraging with high encounter risk and small herd size. Adult females with calves increased scanning and decreased foraging with high encounter risk, but showed no decrease in scanning with large herd size. Yearlings increased scanning and decreased feeding with small herd size, but not with high encounter risk. Adult males were least vigilant, fed most and were not influenced by encounter risk or herd size. These age-sex class differences led to significant differences in group vigilance depending on the composition of the herd. Herds with a majority of mothers were significantly more vigilant than herds with a majority of adult males. However, these differences in group vigilance had no influence on the individual scanning of females without calves. Thus, the decrease in individual scanning with herd size may depend more on changes in individual risk than on cooperative detection of predators. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

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’.     

Vigilance in Przewalski's gazelle: effects of sex, predation risk and group size

Vigilance in social animals is often aimed at detecting predators. Many social and environmental factors influence vigilance, including sex, predation risk and group size. During the summer of 2007, we studied Przewalski's gazelle Procapra przewalskii , an endemic ungulate to the Qinghai-Tibet Plateau, to test whether and how these three factors affect vigilance. We distinguished groups consisting of males, mothers with lambs and females without lambs making observations on groups in the presence or absence of nearby predators. We assessed the group-size effect on vigilance and how this varied with levels of predation risk and sex. Males and mothers scanned longer and with a higher frequency than females without lambs. Individuals were more vigilant under direct predation threat. Although vigilance generally decreased with group size, the extent of the decrease was independent of predation risk and was not significant in males. The results suggest that mothers are more vigilant suggesting greater vulnerability and that males may have increased their vigilance to compete for higher social ranks. The positive correlation between vigilance and predation risk and the negative correlation between vigilance and group size are consistent with earlier findings, but we failed to find an interaction between group size and predation risk on vigilance perhaps because vigilance levels are low even in small groups, thus making similar vigilant upward adjustments in both small and large groups.     

Correlation of relative muzzle width and relative incisor width with dietary preference in ungulates

Qualitative observations suggest that grazing ungulates have relatively broader muzzles than browsing ones, and that grazers have incisors that are all of a similar size, as opposed to the large central and smaller lateral incisors seen in browsers. These differences may be correlated respectively with the need for grazing ungulates to maintain a large daily intake, or for browsing ungulates to forage selectively in a stand of vegetation. Quantitative examination of relative muzzle width and incisor width ratio in 95 species of living ungulates, correlated with seven different types of dietary preferences, substantiated these observations, although phylogenetic history may exhort a strong influence on morphological proportions. For example, equids have relatively narrower muzzles than grazing ruminants despite their less selective mode of feeding. The narrowest relative muzzle widths are not found in regular browsers, but in high level browsers and in mixed feeders in open habitats. Incisor width ratio can distinguish grazers from browsers, but can not be used to discriminate mixed feeders from other feeding types, and grazers appear to have incisors that are relatively broader overall than those of other dietary types, in correlation with their relatively broader muzzles.     

Foraging efficiency and vigilance behaviour of impala: the influence of herd size and neighbour density

Group foraging can be beneficial for ungulates by decreasing the time required for vigilance, but it can also prove costly because of competition. To determine responses to gregarious behaviour, we studied foraging activity and vigilance of impala ( Aepyceros melampus ) near Kruger National Park, South Africa. We measured time spent foraging, vigilant, moving, grooming, engaging in social interactions and determined herd size and group distribution (i.e. density). We calculated accepted food abundance (AFA), food ingestion rate, steps per minute and percent vigilance for female, bachelor male and herd male impala. There was no relationship between herd size and vigilance, but vigilance decreased with increasing density ( t _1,311 = 4.91, P  <0.0001). Additionally, AFA decreased ( t _1,61 = 5.96, P  <0.0001) and steps per minute increased ( t _1,311 = 14.38, P  <   0.0001) as more individuals fed in close proximity to each other. Impala could be altering their behaviour to accommodate a perceived change in resources because of intraspecific competition and these adjustments might be related more to the distribution of herd members than to herd size. Further studies should examine the behaviour of gregarious animals in relation to the distribution of herd members in addition to 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.     

Perch exposure and predation risk: a comparative study in passerines

Singing birds optimize signal transmission by perching in exposed locations. However, conspicuous singing may be risky, and previous research has found that individuals trade off singing performance with song perch exposure. We studied the relationship between predation risk (degree of concealment, height in tree or shrub, and distance to the forest edge) and time allocated to singing and vigilance in a group of 13 passerine species living in an East African savanna. Concealed birds sang more and were less vigilant. Vigilance increased as distance to the forested edge increased, but distance had no effect on time allocated to singing. Body size was significantly correlated with vigilance but not singing; larger passerines were more sensitive to both relative concealment and the distance to the forest edge, while song was influenced by neither of these factors. Perch height had no effect on either behavior. Our results suggest that birds modify vigilance and, to some extent, singing behavior to minimize their exposure to predators.     

Visibility, group size, vigilance, and drinking behavior in coati (Nasua narica) and white-faced capuchins (Cebus capucinus): experimental evidence

I studied the role of visibility on the vigilance behavior of coatis Nasua narica and white-faced capuchins Cebus capucinus in a dry tropical forest in Costa Rica. Vigilance and drinking behavior were observed at a waterhole where the water surface was 2–3 m below ground level for 5 days. Then I provided an artificial water pan of similar size adjacent to the natural waterhole, but level with the ground, and observed the same behaviors. Animals drinking at the natural waterhole had to descend below ground where they could not see above ground, while those drinking from the experimental pan could see approaching predators. Variations in drinking and vigilance behavior were accounted for by site (natural vs. experimental water source) and group size, with site contributing most significantly to variations. Both coatis and capuchins decreased their time to wait at the forest edge before emerging to drink, had more drinking bouts, had longer bout lengths, and decreased their vigilance behavior when they drank at the ground-level waterhole compared to the one below the surface, where their visibility was obscured. For both species, total drinking time increased, and vigilance decreased, with group size. This study demonstrated experimentally that visibility and group size influence drinking and vigilance behaviors. Received: 9 February 2000 / Received in revised form: 3 November 2000 / Accepted: 10 November 2000     

The functional significance of the browser-grazer dichotomy in African ruminants

The allometric relationships for the fermentation rate of dry matter, the total energy concentration of volatile fatty acids (VFAs), the energy supplied from VFA production and the mass of the digesta contents within the rumen or caecum and proximal colon (hindgut) were used to test whether the digestive strategies of grazing and browsing African ruminants differ. The wet and dry mass of the contents of the rumen and hindgut were allometrically related to body mass (BM). These relationships did not differ between browsing and grazing ruminants. The fermentation rates in the rumen were strongly allometric and the intercepts of the relationships did not differ between browsers and grazers. The fermentation rates in the hindgut were not allometrically related to BM and did not differ between ruminants with different feeding habits. Likewise, the total energy concentration of the VFAs in the rumen and hindgut showed no allometric scaling and did not differ between browsing and grazing ruminants. The energy supplied by VFA production in both the rumen and hindgut of African ruminants scaled at around 0.8 with BM. Only in the case of the energy supplied by VFAs in the rumen were there significantly different intercepts for browsing and grazing ruminants. The energy supplied by VFA production in the rumen was inadequate to meet the energy requirements for maintenance of browsers and small grazers. The retention time of digesta in the alimentary tract was positively related to BM although there was no difference in the allometric relationships for grazers and browsers. The results of these analyses suggest that, after controlling for the effects of body mass, there is little difference in digestive strategy between African ruminants with different morphological adaptations of the gut.     

Differences in selective reticulo-ruminal particle retention as a key factor in ruminant diversification

The measurement of passage rate is important for the concept of ruminant diversification. While supporters of Hofmann's 1989 feeding type classification claim that browsing ruminants have faster passage rates than grazing ruminants, other researchers consider the passage rate to depend on body size alone. To date, no convincing comparison of ruminant passage rates has been put forward. For comparative purposes, we suggest the use of the "selectivity factor", which is an expression of how much longer particles of a defined size (<2 mm) are retained in the ruminant digestive tract than fluids. From the limited data available, it seems that grazing ruminants display selectivity factors between 1.56 and 3.80, whereas browsers have a much narrower range of 1.14-1.80. This suggests that browsers are not able to selectively retain particles as long as grazers. Intake of browsers, on the other hand, may not be limited by physical fill of the forestomach to the same degree as in grazers. This result can explain several observations on the digestive physiology of browsers, some of which have been linked to a rumen bypass mechanism. We propose that the ability for selective particle retention is a key factor for understanding the physiological consequences of ruminant diversification.