Alarm Calls Affect Foraging Behavior in Eastern Chipmunks (Tamias striatus, Rodentia: Sciuridae)

We used playback experiments to test whether alarm calls affected the foraging behavior of eastern chipmunks (Tamias striatus). We subjected chipmunks, foraging at artificial feeding stations, to three playback treatments (silent, control noise, and alarm call) and examined changes in vigilant and foraging behavior. Chipmunks responded to alarm calls with a greater degree and duration of vigilant behavior, such as look‐ups and alert postures. Chipmunks also ran a shorter distance to cover and took longer to re‐emerge from the burrow after hearing an alarm call. Alarm calls caused individuals to spend more time exposed at the feeding stations; however, individuals also took significantly fewer seeds after hearing an alarm call. This was not due to a difference in the time spent handling food, but rather to a slower rate of loading. Chipmunks appear to sacrifice energy gain by increasing vigilance after hearing an alarm call. This study suggests that to avoid the costs of unnecessary escape behavior, individuals directly assess their own risk rather than relying only on indirect cues such as alarm calls.     

Functions of Intermittent Locomotion in Mustached Tamarins (<Emphasis Type="Italic">Saguinus mystax</Emphasis>)

Many animals interrupt their moving with brief pauses, which appear to serve several different functions. We examined the function of such intermittent locomotion in wild living mustached tamarins (Saguinus mystax), small arboreal New World primates that form mixed-species groups with saddleback tamarins (Saguinus fuscicollis). We investigated how different environmental and social factors affect pausing during locomotion and used these data to infer the function of this behavior. As measures of intermittent locomotion, we used percentage of time spent pausing and pause rate. We considered 3 possible functions that are not mutually exclusive: increased endurance, route planning, and antipredator vigilance. Mustached tamarins spent on average (mean ± SE) 55.1 ± 1.0% of time pausing, which makes effective resource exploitation more time consuming and needs to be outweighed by correspondingly large benefits. Percentage of time spent pausing decreased in larger mixed-species groups vs. smaller mixed-species groups and decreased with height and in monkeys carrying infants. It was not affected by sex, age, spatial arrangement, or single-species group size. Pause rate increased in individuals traveling independently compared to those traveling in file, but was not affected by other factors. The group size effect in mixed-species groups lends support to the notion that pausing during locomotion is an antipredator tactic that can be reduced in the increased safety of larger groups, but other results suggest that additional functions, particularly route planning, are also of great importance. Benefits in terms of predator confusion and group movement coordination are also likely to play a role and remain a topic for further research.     

Load Size Variation in the Eastern Chipmunk,Tamias striatus: The Importance of Distance from Burrow and Canopy Cover

We examined the degree to which chipmunks foraging from dishes containing sunflower seeds varied cheek pouch load size according to both travel time (distance from burrows) and predation hazard (canopy cover). Results showed that larger loads were taken under cover at some distance from burrows than were taken from closer or more open sites. Distance effects appeared to reflect the influence of traveling times on quitting harvest rates as predicted by central place foraging models. Quitting harvest rates also appeared to be higher under low than under dense canopy cover; this trend was most pronounced at some distance from burrows suggesting that foraging decisions were made relative to both energetics and predator avoidance. Chipmunks appeared to employ a patch exploitation strategy that minimized the hazard:gain ratio.     

Linking summer foraging to winter survival in yellow pine chipmunks (Tamias amoenus)

This study links summer foraging and scatter-hoarding to winter larder-hoarding and winter survival in yellow pine chipmunks (Tamias amoenus) by comparing patterns of time allocation and winter larder contents in 2 years with very different levels of resource availability. In 2003, seed production and the number of trees and shrubs producing seeds were high. In 2004 seed crops were small. Chipmunks allocated more time to foraging when food resources were scarce (66% in 2004) compared to when they were abundant (39% in 2003). Increased time allocated to foraging in 2004 corresponded to significant decreases in time allocated to vigilance, resting, and social interactions. When seeds were scarce (i.e., in autumn 2004), chipmunks spent more time searching for cached food items than gathering seeds from plants or the ground surface. Despite the increase in foraging effort, the edible mass and caloric contents of larders were significantly smaller in 2004. In the year with low seed production, the diversity of seed species found in larders increased, and many of these seeds were of species that ripened in summer. When autumnal seed production by Jeffrey pine seeds was high, Jeffrey pine seeds were nearly the exclusive food item found in larders. Larder contents would have provisioned chipmunks for an estimated 116-257 days in 2003 and but only 6-111 days in 2004. It is likely that all chipmunks would have survived the winter of 2003 (duration ~110-120 days). However, none of the larders recovered in 2004 contained enough food to have provisioned the inhabitant for the ~148-158 days of winter.     

Interference competition in central place foragers: the effect of imposed waiting on patch-use decisions of eastern chipmunks, Tamias striatus

When central place foragers compete aggressively for patchyresources, subordinates may be preventedfrom collecting fooduntil a dominant has departed with its load. Extensions of centralplace foraging models predict that animals forced to wait ata patch should increase their load sizes and patch times aswellas their tendency to search for and switch to alternative patches.We tested these predictions usingeastern chipmunks, Tamias striatus,hoarding sunflower seeds collected from seed/vermiculite mixturesintrays placed 5-8 m from their burrows. By using her hand toprevent access to the patch, the experimentersubjected animalsto progressively increasing waiting times at two seed densities;another series of trialsat the same seed densities monitoreda similar number of trips without imposed waiting. As predicted,patch times and load sizes were higher in sessions with imposedwaiting than in control sessions. Loadsizes increased with trialnumber in experimental sessions but decreased or remained thesame in controlsessions. Chipmunks spent more of their timesearching for alternative patches during trials with imposedwaiting than during controls. They also started searching foralternative patches at lower levels of imposed waiting whenusing poor than when using rich patches. These results indicatethat the effects of interference on foraging decisions and onspatial overlap between individuals can be predicted by simpleeconomic models. Furthermore, the results suggest how resource-defensetactics can be predicted by the economic effects of interferenceon the foraging efficiency of the opponent.     

The influence of habitat on travel speed, intermittent locomotion, and vigilance in a diurnal rodent

We studied effects of habitat structure on routine travel velocities, intermittent locomotion, and vigilance by the degu (Octodondegus), a diurnal rodent of central Chile. We predicted thattravel speed, pauses during locomotion, and vigilance wouldbe greater in open (riskier) than in shrub (safer) habitats.Video recordings of marked individuals in the wild were used to measure speed and other variables of spontaneous locomotionnot triggered by predatory attack or any other noticeable stimulusduring nonforaging periods. Time spent vigilant while foragingwas also measured. Because degus use bare-ground runways fordistant movements (e.g., between burrow openings and/or foodpatches), data on locomotion decisions were not confounded by effects of obstructive vegetation cover and/or resource abundance.When moving across the habitat between different feeding places,degus showed an intermittent pattern of locomotion, interruptingrunning events with short pauses. As predicted, travel speedand the duration of pauses between locomotion bursts were significantlygreater in open habitats. Further, the duration of locomotionbursts between feeding sites or between feeding sites and burrowswas significantly longer in open habitats. Our assumption that pauses and velocities are independent decisions was supportedby the lack of correlation between pauses and speeds duringlocomotion events. During foraging, degus devoted more timeto vigilance in open than in shrub habitats. The static positionadopted by degus during pauses, the speeds attained during movements, and the concordance between pausing behavior andvigilance across habitats suggest that pausing has an antipredatoryrole and is not limited to orientation and/or physiologicalrecovery. Our results support the view that degus perceivehigher predation risk in open areas and that flexible movement behavior reflects an adaptive antipredator response.     

Switching to a feeding method that obstructs vision increases head-up vigilance in dabbling ducks

A major assumption in most models of foraging is that feeding and vigilance are mutually exclusive. A recent experimental study challenged this hypothesis and demonstrated that birds are able to detect predators while pecking seeds on the ground (head-down vigilance). Experimental obstruction of head-down vigilance makes birds increase head-up vigilance (i.e. the classical overt vigilance posture). For many foragers in the wild, visibility varies between habitats and foraging methods. We compared the vigilance of Teal Anas crecca and Shoveler Anas clypeata when foraging with their eyes above the water surface (shallow feeding, only the bill submerged) and when foraging with their eyes underwater (deep feeding, head and neck underwater, or upending), at three wintering sites in western France. Birds of both species spent less time in head-up vigilance during shallow foraging than during deep foraging, with no significant difference between sites, which suggests that they are capable of some vigilance during shallow foraging. During deep foraging, the time spent vigilant increased because the frequency of scans was much higher, while scan length decreased. However, these differences could have resulted from variations in the availability of food at different depths. In an experiment where the food availability was constant, we observed the same pattern, with a higher frequency of scans during deep foraging. This study therefore provides strong support for the idea that vigilance and foraging are not always mutually exclusive and shows that switching between searching methods can cause vigilance time – and, as a consequence, loss of feeding time – to vary. This should be taken into account in future field and experimental studies of the trade-off animals make between vigilance and feeding.     

Daily routines and predator encounters in Yellowhammers Emberiza citrinella in the field during winter

Diurnal and seasonal variation in foraging behaviour of Yellowhammers Emberiza citrinella and activity of their avian predators were studied in an unmanipulated winter field situation around Uppsala, Sweden. In December, when time available for foraging was lowest, Yellowhammers seemed to be time-stressed. In order to meet their energetic needs, they reduced the time allocated to vigilance and increased the time allocated to foraging. Yellowhammers did not systematically change the time spent foraging during the day in December, which indicates time-stress, while they decreased the time spent foraging during the day in both November and February. Predator activity was highest in the afternoon, when Yellowhammers spent the least time foraging. Yellowhammers may have adopted a routine with decreasing foraging rates over the day in November and February, when time available for foraging was longer, in order to avoid foraging during periods of high predator activity. The diurnal activity pattern of predators together with daylength and energetic needs are factors that might be important in shaping daily foraging routines in small birds.     

Information transfer and the facilitation and inhibition of feeding in a schooling fish

Synopsis Recent studies show that fish forage actively when perceived risk is low, but decrease foraging and increase vigilance when perceived risk is high. Isolated juvenile chum salmon,Oncorhynchus keta, were visually exposed to groups of conspecifics engaged in different activities to examine their ability to gain information about foraging opportunity and risk by interpreting conspecific behavior. Isolates ate most when exposed to feeding groups, less when exposed to nonfeedig groups, and least when exposed to alarmed groups. Isolates exposed to alarmed conspecifics also spent more time motionless than did fish exposed to either feeding or nonfeeding conspecifics. These findings indicate that schooling fish gain information by interpreting conspecific behavior, and are consistent with research showing that animals balance the conflicting demands of foraging and vigilance.     

东方田鼠家族群对成员个体觅食行为的影响

植食性哺乳动物在分享社群觅食带来好处的同时,是否因个体间的相互干扰而影响其摄入率。在新鲜马唐叶片构建的均质密集食物斑块上,测定东方田鼠家族群成员个体在食物斑块上的觅食行为序列过程及行为参数,检验家族群存在对成员个体觅食行为的影响。结果发现,东方田鼠家族群雌、雄成员个体的觅食行为参数均无显著差异。然而,与单只个体相比,家族群觅食尽管能显著地缩短成员个体的觅食决定时间,但却显著地降低了成员个体的摄入率。分析觅食行为参数觅食中断时间发现,相较于单只个体,家族群成员个体间因相互干扰而引起的觅食中断时间的增加,不但增大了收获每口食物的时间,而且导致其摄入率下降。检测家族群成员个体各警觉行为动作参数,发现,成员个体间的相互干扰能引致个体的一般扫视、盯视及嗅闻动作时间比例显著增大,尽管直立扫视和静听动作时间比例减少显著,但并未使个体的觅食中断时间减小。结果充分说明,东方田鼠家族群成员个体间的相互干扰能使个体觅食行为参数发生变异,导致觅食中断时间增加,摄入率降低。     

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.     

How to climb a tree: lizards accelerate faster,but pause more,when escaping on vertical surfaces

Many species of lizards effectively traverse both two and three‐dimensional habitats. However, few studies have examined maximum locomotor performance on different inclines. Do maximum acceleration and velocity differ on a level and inclined surface? Do lizards pause more on an inclined surface? To address these questions, Sceloporus woodi lizards (N = 12) were run in the laboratory on a level trackway and a vertical tree trunk. This species is known to frequently utilize both vertical and horizontal aspects of its habitat. Average maximum acceleration on the vertical surface exceeded that on the level surface, although average maximum velocity exhibited the opposite pattern. The average number of pauses during level locomotion was lower compared to vertical locomotion. In addition, the average location of the first pause on the level surface was 0.51 m, which is farther than the average for vertical locomotion where the first pause was at 0.35 m. The combination of performance and pause data suggests that the relative lack of pausing during level locomotion allows individuals to reach higher maximum velocities on level surfaces because they accelerate over greater distances. The increased pausing when moving vertically could be a result of high energetic demands of vertical locomotion, or greater microhabitat complexity as a result of branching and/or refuges. The faster acceleration exhibited during vertical locomotion by S. woodi likely offsets the frequent pauses. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 83–90.     

Saltatory Search in Free-Living <Emphasis Type="Italic">Callithrix jacchus</Emphasis>: Environmental and Age Influences

Researchers generally categorize motile foraging behavior into 3 strategies: ambush, cruise, and saltatory searching. During saltatory searches, predators move from one location to the next, frequently pausing to scan for prey that are hard to detect and widely distributed. We investigated whether 1) the foraging strategy of free-living common marmosets conforms to the strategy; 2) scanning occurs solely when the individual is stationary; 3) the environment (dense and sparse vegetation) influences foraging behavior; and 4) the age of the marmosets is related to their foraging behavior. Bezerra carried out the observations in a 32-ha fragment of Atlantic Forest in the Northeast of Brazil and in an adjacent condominium. Using the focal sampling method, we observed 31 common marmosets, including adults, juveniles, and infants, Bezerra recorded the following behaviors (in dense and sparse vegetation): locomotion (subdivided into minor locomotion—moving distances ≤1 m; major locomotion—moving distances >1 m), scan, pause, and capture. The frequency of scanning behavior was significantly greater when individuals were stationary. Adults and juveniles exhibited the clearest differentiation in terms of locomotion, both adjusting the behavior in accordance with the environment; periods of major locomotion were more frequent in sparse vegetation, whereas minor locomotion was more frequent in dense vegetation. In contrast, infants exhibited major locomotion more frequently in dense vegetation. We conclude that common marmosets use a saltatory strategy when foraging, and that their foraging behavior is plastic, changing both with the age of the individual and with the density of the vegetation.     

Sexual,Seasonal and Group Size Differences in the Allocation of Time between Vigilance and Feeding in the Greater Rhea,Rhea americana

We studied the effect of sex and group size on the proportion of time a greater rhea, Rhea americana, allocates to vigilance and feeding during the breeding and the non-breeding seasons. We analysed 175 records of focal animals that were feeding alone or in groups of 2 to 26 birds. In both seasons, males spent more time in vigilance and less time in feeding than females. Both sexes spent more time in vigilance and less time in feeding during the breeding season. Sexual and seasonal differences in vigilance were the result of different mechanisms. Males had shorter feeding bouts than females but there were no sexual differences in the length of the vigilance bouts. On the contrary, seasonal differences were the result of males and females having longer vigilance bouts during the breeding season but there were no seasonal differences in the length of the feeding bouts. During the non-breeding season, individual vigilance was higher in rheas foraging alone than in groups. In this case, solitary birds had longer vigilance and shorter feeding bouts than birds foraging in groups. We discuss the possible effect of intragroup competition and food availability on the allocation of time between feeding and vigilance in this species.     

Vigilance during food handling by Oystercatchers Haematopus ostralegus reduces the chances of losing prey to kleptoparasites

Oystercatchers Haematopus ostralegus periodically pause while handling mussels Mytilus edulis to make visual scans. This paper presents evidence that scanning is associated with the high incidence of intra-specific food stealing among mussel-eating Oystercatchers. Scanning increased in frequency as bird density - and the risk of being attacked for mussels increased and the duration of attacks decreased. Additionally, among a sample of individually marked adults, the aggressive dominant birds spent half as much time scanning as the less aggressive subdominants and were also less likely to be attacked. Whereas detecting an attack made no difference to the success with which the dominants defended their mussels, subdominants increased their chances of retaining the mussel if they detected and carried the mussel away from the approaching attacker. The extra time which the less aggressive birds spent in vigilance seems best understood as a tactic for reducing food loss to kleptoparasites.     

Sex differences in giraffe foraging behavior at two spatial scales

We test predictions about differences in the foraging behaviors of male and female giraffes (Giraffa camelopardalis tippelskirchi Matchie) that derive from a hypothesis linking sexual size dimorphism to foraging behavior. This body-size hypothesis predicts that males will exhibit specific behaviors that increase their dry-matter intake rate relative to females. Foraging behavior was examined at two hierarchical levels corresponding to two spatial and temporal scales, within patches and within habitats. Patches are defined as individual trees or shrubs and habitats are defined as collections of patches within plant communities. Males were predicted to increase dry-matter intake rate within patches by taking larger bites, cropping bites more quickly, chewing less, and chewing faster. Within habitats, males were expected to increase intake rate by increasing the proportion of foraging time devoted to food ingestion as opposed to inter-patch travel time and vigilance. The predictions were tested in a free-ranging population of giraffes in Mikumi National Park, Tanzania. Males spent less total time foraging than females but allocated a greater proportion of their foraging time to forage ingestion as opposed to travel between patches. There was no sex difference in rumination time but males spent more time in activities other than foraging and rumination, such as walking. Within patches, males took larger bites than females, but females cropped bites more quickly and chewed faster. Males had longer per-bite handling times than females but had shorter handling times per gram of intake. Within habitats, males had longer average patch residence times but there was no significant sex difference in inter-patch travel times. There was no overall difference between sexes in vigilance while foraging, although there were significant sex by habitat and sex by season interactions. Although not all the predictions were confirmed, overall the results agree qualitatively with the body-size hypothesis. Sex-related differences in foraging behavior led to greater estimated intake rates for males at the within-patch and within-habitat scales. Received: 20 November 1995 / Accepted: 5 November 1996     

The effect of group size on vigilance in Ruddy Turnstones Arenaria interpres varies with foraging habitat

Foraging birds can manage time spent vigilant for predators by forming groups of various sizes. However, group size alone will not always reliably determine the optimal level of vigilance. For example, variation in predation risk or food quality between patches may also be influential. In a field setting, we assessed how simultaneous variation in predation risk and intake rate affects the relationship between vigilance and group size in foraging Ruddy Turnstones Arenaria interpres. We compared vigilance, measured as the number of ‘head‐ups’ per unit time, in habitat types that differed greatly in prey energy content and proximity to cover from which predators could launch surprise attacks. Habitats closer to predator cover provided foragers with much higher potential net energy intake rates than habitats further from cover. Foragers formed larger and denser flocks on habitats closer to cover. Individual vigilance of foragers in all habitats declined with increasing flock size and increased with flock density. However, vigilance by foragers on habitats closer to cover was always higher for a given flock size than vigilance by foragers on habitats further from cover, and habitat remained an important predictor of vigilance in models including a range of potential confounding variables. Our results suggest that foraging Ruddy Turnstones can simultaneously assess information on group size and the general likelihood of predator attack when determining their vigilance contribution.     

鄱阳湖围垦区藕塘越冬白鹤的时间分配与行为节律

2016年12月—2017年3月,采用瞬时扫描法研究了鄱阳湖围垦区藕塘生境中白鹤Grus leucogeranus越冬期的行为,共扫描2560次,23219只次,包括18031只次成鹤和5188只次幼鹤。结果表明,觅食(41.78%)、警戒(25.02%)、修整(15.00%)和休息(10.84%)是白鹤越冬期的主要行为。与自然生境相比,藕塘生境中白鹤主要采取多休息和修整的策略来节省能量支出。成鹤的觅食行为(35.29%)显著低于幼鹤(62.42%)(F_(1,12)=45.977,P0.01),警戒行为(28.66%)则显著高于幼鹤(10.26%)(F_(1,12)=38.975,P0.01)。家庭群成鹤觅食行为(43.96%)极显著高于非家庭群成鹤(27.04%)(F_(1,12)=60.169,P0.01)。家庭群成鹤需要喂食幼鹤,它们花费更多的时间觅食弥补能量的消耗。行为节律上,白鹤各时段觅食行为占总行为的比例均较高,11:00—11:59出现明显高峰,占总行为的48.64%。警戒行为无明显的低谷和高峰。成鹤在各时段的觅食行为比例均明显低于幼鹤,成鹤和幼鹤的觅食曲线变化趋势相似,即觅食比例的升高和降低比较同步。成鹤在各时段的警戒行为和修整行为比例均明显高于幼鹤,成幼鹤的休息行为比例在各时段差别不大,均在14:00—14:59有个明显的高峰。家庭群成鹤的觅食行为比例几乎在各时段均明显高于非家庭群,家庭群成鹤的修整行为和休息行为在大部分时段均低于非家庭群成鹤。因此家庭群成鹤采取多取食,少休息和修整的策略提高自身的适合度,同时保证对后代的抚育。     

Effect of Group Size on Individual and Collective Vigilance in Greater Rheas

We studied the effect of group size on the proportion of time that greater rheas, Rhea americana, allocated to vigilance and feeding during the non‐breeding season. We tested whether: (1) the proportion of time that one bird allocates to vigilance (individual vigilance) decreases with group size, and (2) the proportion of time that at least one bird of the group is vigilant (collective vigilance) increases with group size. We analyzed video‐recordings of birds that were foraging alone or in groups from two to 12 birds. The proportion of time allocated to individual vigilance decreased and the proportion of time spent feeding increased with group size. In both cases the main significant difference was between birds foraging alone and in groups. Collective vigilance did not vary with group size and it was lower than expected if vigilance bouts were random or sequential. Our results indicate that rheas foraging in large groups would not receive the benefit of an increase in collective vigilance, although they could still benefit from a reduction of predation risk by the dilution effect.     

Spatial Heterogeneity and Nestmate Encounters Affect Locomotion and Foraging Success in the Ant Dorymyrmex goetschi

The spatial structure of habitats contains physical barriers that restrict the performance of diverse behavioural tasks. In heterogeneous habitats, information acquisition may allow animals to improve the performance of diverse activities such as foraging and locomotion. Nonetheless, changes in locomotion performance and their effects on the foraging success of animals have been scarcely studied. We examined these relationships in the harvester ant Dorymyrmex goetschi (subfamily Dolichoderinae) under laboratory conditions. In an experimental arena, we offered a food patch located at a fixed distance from the nest entrance. Landscape heterogeneity was created using wooden cubes arranged in different types of spatial distribution. We video recorded the behaviour of different colonies and quantified the number of active foragers, number of head contacts per capita per inbound trip, path length by workers that transported a food load from the resource patch to the nest, time invested in inbound travels, and the number of prey captured per colony. During the initial phase of patch exploitation, the number of foragers and prey captured were significantly lower than during the half and final phases of the experiment. Landscapes with greater spatial heterogeneity increased travel time and diminished locomotion velocity. A multiple regression analysis revealed that greater antennal contacts and locomotion velocities increased prey removal. Therefore, in this study, we documented a formal link between variables that characterize the movement paths of individuals and the foraging success of a colony. Information transfer between individuals generated a collective work with a concomitant improvement of food exploitation.