视野受阻对东方田鼠觅食行为的影响

植食性小型哺乳动物的觅食行为是对特定生境的适应性产物。食物斑块周边植被对动物视野遮挡是否通过作用于其觅食活动中的警觉而影响摄入率。采用新鲜白三叶叶片构建东方田鼠密集均质食物斑块,以牛皮纸模拟食物斑块周边植被遮挡田鼠视野,测定其在食物斑块上的觅食行为序列过程及行为参数,检验食物斑块周边植被高度对东方田鼠摄入率的影响。结果发现,个体在不同程度视野受阻条件下食物摄入率无显著差异。分析觅食行为参数动态发现,在不同视野受阻条件下,个体能通过调整各采食回合内警觉行为动作的发生频次和持续时间,维持觅食回合内总的觅食中断时间的稳定,进而保证进食时间的稳定。东方田鼠在不同程度的视野遮挡条件下均能通过行为变异和优化使摄入率保持稳定。结果亦充分说明,东方田鼠在警觉强度上的变化不能反映觅食中断所带来的食物收益减损的代价,但觅食活动中各警觉动作的持续时间的变异却能够明确指示个体摄入率的动态变化,因此,以觅食活动中警觉引起的觅食中断时间代价为线索,检验其对摄入率的影响,是评价植食性小型哺乳动物在不同生境觅食适应性策略的有效的方法。     

The influence of bite size on foraging at larger spatial and temporal scales by mammalian herbivores

Organisms respond to their heterogeneous environment in complex ways at many temporal and spatial scales. Here, I examine how the smallest scale process in foraging by mammalian herbivores, taking a bite, influences plants and herbivores over larger scales. First, because cropping bites competes with chewing them, bite size influences short-term intake rate of herbivores within plant patches. On the other hand, herbivores can chew bites while searching for new ones, thus influencing the time spent vigilant and intake rate as animals move among food patches. Therefore, bite size affects how much time herbivores must spend foraging each day. Because acquiring energy is necessary for fitness, herbivores recognize the importance of bite size and select bites, patches and diets based on tradeoffs between harvesting rates, digestion, and sheering forces. In turn, induced structural defenses of plants, such as thorns, allow plants to respond immediately to herbivory by reducing bite size and thus tissue loss. Over evolutionary time, herbivores have adapted mouth morphology that allows them to maximize bite size on their primary forage plant, whereas plants faced with large mammalian herbivores have adapted structures such as divarication that minimize bite size and protect themselves from herbivory. Finally, bite size available among plant communities can drive habitat segregation and migration of larger herbivores across landscapes.     

Effects of Humans on Behaviour of Wildlife Exceed Those of Natural Predators in a Landscape of Fear

Background

Human disturbance can influence wildlife behaviour, which can have implications for wildlife populations. For example, wildlife may be more vigilant near human disturbance, resulting in decreased forage intake and reduced reproductive success. We measured the effects of human activities compared to predator and other environmental factors on the behaviour of elk (Cervus elaphus Linnaeus 1758) in a human-dominated landscape in Alberta, Canada.

Methodology/Principal Findings

We collected year-round behavioural data of elk across a range of human disturbances. We estimated linear mixed models of elk behaviour and found that human factors (land-use type, traffic and distance from roads) and elk herd size accounted for more than 80% of variability in elk vigilance. Elk decreased their feeding time when closer to roads, and road traffic volumes of at least 1 vehicle every 2 hours induced elk to switch into a more vigilant behavioural mode with a subsequent loss in feeding time. Other environmental factors, thought crucial in shaping vigilance behaviour in elk (natural predators, reproductive status of females), were not important. The highest levels of vigilance were recorded on public lands where hunting and motorized recreational activities were cumulative compared to the national park during summer, which had the lowest levels of vigilance.

Conclusions/Significance

In a human-dominated landscape, effects of human disturbance on elk behaviour exceed those of habitat and natural predators. Humans trigger increased vigilance and decreased foraging in elk. However, it is not just the number of people but also the type of human activity that influences elk behaviour (e.g. hiking vs. hunting). Quantifying the actual fitness costs of human disturbance remains a challenge in field studies but should be a primary focus for future researches. Some species are much more likely to be disturbed by humans than by non-human predators: for these species, quantifying human disturbance may be the highest priority for conservation.     

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.      

Group size effects on foraging and vigilance in migratory Tibetan antelope

Large group sizes have been hypothesized to decrease predation risk and increase food competition. We investigated group size effects on vigilance and foraging behaviour during the migratory period in female Tibetan antelope Pantholops hodgsoni, in the Kekexili Nature Reserve of Qinghai Province, China. During June to August, adult female antelope and yearling females gather in large migratory groups and cross the Qinghai-Tibet highway to calving grounds within the Nature Reserve and return to Qumalai county after calving. Large groups of antelope aggregate in the migratory corridor where they compete for limited food resources and attract the attention of mammalian and avian predators and scavengers. We restricted our sampling to groups of less than 30 antelopes and thus limit our inference accordingly. Focal-animal sampling was used to record the behaviour of the free-ranging antelope except for those with lambs. Tibetan antelope spent more time foraging in larger groups but frequency of foraging bouts was not affected by group size. Conversely, the time spent vigilant and frequency of vigilance bouts decreased with increased group size. We suggest that these results are best explained by competition for food and risk of predation.     

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 habitat complexity on the functional response of a seed-eating passerine

Recent population declines of seed-eating farmland birds have been associated with reduced overwinter survival due to reductions in food supply. An important component of predicting how food shortages will affect animal populations is to measure the functional response, i.e. the relationship between food density and feeding rate, over the range of environmental conditions experienced by foraging animals. Crop stubble fields are an important foraging habitat for many species of seed-eating farmland bird. However, some important questions remain regarding farmland bird foraging behaviour in this habitat, and in particular the effect of stubble on farmland bird functional responses is unknown. We measured the functional responses of a seed-eating passerine, the Chaffinch Fringilla coelebs , consuming seeds placed on the substrate surface in three different treatments: bare soil, low density stubble and high density stubble. Stubble presence significantly reduced feeding rates, but there was no significant difference between the two stubble treatments. Stubble reduced feeding rates by reducing the maximum attack distance, i.e. the distance over which an individual food item is targeted and consumed. The searching speed, handling time per seed, proportion of time spent vigilant, duration of vigilance bouts and duration of head-down search periods were unaffected by the presence of stubble. The frequency of vigilance bouts was higher in the bare soil treatment, but this is likely to be a consequence of the increased feeding rate. We show the influence of a key habitat type on the functional response of a seed-eating passerine, and discuss the consequences of this for farmland bird conservation.     

The influence of snow on the functional response of grazing ungulates

The forage intake rate of grazing ungulates is limited either by the rate at which they encounter food items, or the rate at which food items are handled. Whether an ungulate is encounter‐ or handling‐limited influences spatial and temporal depletion of forage, daily time budgets, and ultimately animal condition. Previously, vegetation abundance has been used as a surrogate for an ungulate's encounter rate with food items and related to observed bite rate to determine whether intake rate is encounter‐ or handling‐limited. In temperate climates snow accumulation during winter limits access to vegetation by forcing animals to wade and paw through snow to consume underlying vegetation, increasing the amount of time required to encounter a food item. As a result, an ungulate may be handling‐limited when foraging in a high biomass system under snow‐free conditions, but becomes encounter‐limited when snow accumulates. We derived a model that provides a frame work for estimating the rate at which a grazing ungulate encounters vegetation by considering foraging velocity, vegetation biomass and the time required to paw away snow when present. We then used data from focal observations of 36 wild elk Cervus canadensis wintering on a montane grassland in the Canadian Rockies of Alberta, Canada, to apply our model and estimate encounter rate over a range of vegetation abundance and snow conditions. Using AIC_c in a model selection approach we found that an asymptotic regression model of observed bite rate as a function of estimated encounter rate provided a better fit than similar models using only vegetation abundance as the explanatory variable. An asymptotic model suggests elk were handling‐limited in the absence of snow, but became encounter‐limited when snow accumulated. Our results demonstrate the importance of considering the influence of factors other than vegetation abundance on the intake rate of grazing ungulates.     

Coordination and synchronisation of anti-predation vigilance in two crane species

Much of the previous research on anti-predation vigilance in groups has assumed independent scanning for threats among group members. Alternative patterns that are based on monitoring the vigilance levels of companions can also be adaptive. Coordination of vigilance, in which foragers avoid scanning at the same time as others, should decrease the odds that no group member is alert. Synchronisation of vigilance implies that individuals are more likely to be vigilant when companions are already vigilant. While synchronisation will increase the odds that no one is vigilant, it may allow a better assessment of potential threats. We investigated temporal sequences of vigilance in family flocks consisting of two parents and at most two juveniles in two species of cranes in coastal China. We established whether the observed probability that at least one parent is alert was greater (coordination) or lower (synchronisation) than that predicted under the null hypothesis of independent vigilance. We documented coordination of vigilance in common cranes (Grus grus) foraging in an area with high potential for disturbance by people. We documented synchronisation of vigilance in red-crowned cranes (Grus japonensis) in the less but not in the more disturbed area. Coordination in small flocks leads to high collective vigilance but low foraging rates that may not be suitable in areas with low disturbance. We also argue that synchronisation should break down in areas with high disturbance because periods with low vigilance are riskier. Results highlight the view that temporal patterns of vigilance can take many forms depending on ecological factors.     

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.     

饥饿和食物单宁酸对东方田鼠(Microtus fortis)食物摄入量和觅食行为的影响

实验室条件下,测定饥饿和食物单宁酸对东方田鼠食物摄入量和觅食行为的影响。结果表明,饥饿使实验个体的食物总摄入量增加,食物摄入率及口量大小随饥饿强度的增大而增加,而觅食频次则无显著改变,实验个体每取食回合的觅食时间呈缓慢增加的趋势,与对照组比较,觅食时间差异不显著。东方田鼠优先选择0%单宁酸食物,次为3%单宁酸食物,而对6%单宁酸食物的摄入量最少。在饥饿条件下,东方田鼠食物摄入率的增加主要源于其口量大小,觅食频次和觅摄食时间对食物摄入量增加的贡献不显著。在饥饿条件下,植食性小哺乳动物并未通过延长觅食时间,降低用于防卫、繁殖活动时间来增加食物摄入量,而是通过增加口量大小,提高其食物摄入率来满足其营养需要。验证了饥饿与植物次生化合物共同作用引起田鼠类动物生理的改变,能影响其食物摄入量及觅食行为的假设。     

Antipredator vigilance of juvenile and adult thirteen-lined ground squirrels and the role of nutritional need

Juvenile thirteen-lined ground squirrels, Spermophilus tridecemlineatus, are less vigilant (i.e. they spend less time visually scanning the environment) than adults. To determine whether nutritional need was a potential cause of this difference, we supplemented two groups of free-ranging juveniles during the predispersal stage, while juveniles were still near and around the natal burrows. The high-energy food group (HEF: 11 squirrels) received peanut butter and oats while the low-energy food group (LEF: seven squirrels) received lettuce. Adults (14 squirrels) were also supplemented, but due to their greater home range sizes, it was not feasible to classify them as either HEF or LEF. To evaluate the effect of supplementation on antipredator vigilance, the behavioural act of visually scanning for predators, we videotaped individuals while they were foraging above ground during 5-min observation periods. Each squirrel was observed and weighed during three time periods over 23 days. From the videotape, we extracted measures of time spent vigilant, locomoting and foraging. All three categories of squirrels gained mass over the study period, but the HEF juveniles rapidly exceeded that of the LEF juveniles. Early in the study, LEF and HEF juveniles did not significantly differ in either body mass or time budgets, and, initially, both juvenile groups were similar to adults in the amount of time devoted to vigilance. Later in the study, the behaviour of HEF juveniles closely resembled that of adults (increased time devoted to vigilance and decreased time devoted to foraging), while LEF juveniles decreased vigilance and increased their foraging time. This study indicates that for thirteen-lined ground squirrels the lower vigilance of juveniles is due, at least in part, to the greater nutritional needs of young animals with consequent increases in foraging, which is largely incompatible with vigilance. Copyright 2000 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’.     

To scan or not to scan? Occurrence of the group‐size effect in a seasonally nongregarious forager

Group‐living requires a compromise between safety and direct/indirect costs for individuals. The larger is the group, the greater is the collective vigilance, leading to a greater net food intake per forager because of the time saved individually from scanning behaviour. In turn, individual alertness usually decreases with increasing group size (“group‐size effect”). Information on the occurrence of group‐size effect is still unclear. Previous studies have shown that it may fail to occur or even reverse, for example when costs of interference between conspecifics are high. In turn, assessing whether the group‐size effect would occur in weakly or seasonally gregarious species may help to understand its drivers. We evaluated the occurrence and the extent of group‐size effect in a seasonally nongregarious herbivore, the roe deer Capreolus capreolus. We examined the roles of sex/age class and season as drivers of vigilance behaviour. In roe deer, the group‐size effect did not depend on sex/age class: time spent foraging increased with increasing group size; in turn, vigilance increased with decreasing group size, in all sex/age classes. Females with fawns were the most vigilant sex/age class, thus revealing the cost of offspring protection. Accordingly, the higher spring vigilance levels of does could be related to reproductive costs (e.g., defence of newborn fawns). Conversely, the greater summer vigilance of bucks could result from patrolling/defence of territories. Both adult males and females also showed the higher vigilance in winter, likely because of an increase in the perception of predation risk and/or, possibly, hormones linked to an increase in intolerance of conspecifics, in males. However, the group‐size effect occurred in all the seasons, for adult males and females. Our findings suggest that foraging benefits provided by the group‐size effect may have overcome costs of group‐living, even in a weakly gregarious forager.     

Risky foraging leads to cost-free mate guarding in male teal Anas crecca

Mate guarding by males is common in species with long-lasting pair bonds. We tested if the need to guard females affected foraging depth in male teal (Anas crecca), and if they were more vigilant than females when foraging with submerged eyes (preventing monitoring of competing males and predators). These predictions were not supported, suggesting that foraging depth selection is primarily driven by other factors, presumably food related. A likely reason why deeply foraging males did not increase vigilance is that 37.5% of the foraging time was already dedicated to it. The apparent lack of guarding costs in foraging male teal may explain why such small ducks can maintain pair bonds for up to 7 months.     

Longer guts and higher food quality increase energy intake in migratory swans

1. Within the broad field of optimal foraging, it is increasingly acknowledged that animals often face digestive constraints rather than constraints on rates of food collection. This therefore calls for a formalization of how animals could optimize food absorption rates. 2. Here we generate predictions from a simple graphical optimal digestion model for foragers that aim to maximize their (true) metabolizable food intake over total time (i.e. including nonforaging bouts) under a digestive constraint. 3. The model predicts that such foragers should maintain a constant food retention time, even if gut length or food quality changes. For phenotypically flexible foragers, which are able to change the size of their digestive machinery, this means that an increase in gut length should go hand in hand with an increase in gross intake rate. It also means that better quality food should be digested more efficiently. 4. These latter two predictions are tested in a large avian long-distance migrant, the Bewick's swan (Cygnus columbianus bewickii), feeding on grasslands in its Dutch wintering quarters. 5. Throughout winter, free-ranging Bewick's swans, growing a longer gut and experiencing improved food quality, increased their gross intake rate (i.e. bite rate) and showed a higher digestive efficiency. These responses were in accordance with the model and suggest maintenance of a constant food retention time. 6. These changes doubled the birds' absorption rate. Had only food quality changed (and not gut length), then absorption rate would have increased by only 67%; absorption rate would have increased by only 17% had only gut length changed (and not food quality). 7. The prediction that gross intake rate should go up with gut length parallels the mechanism included in some proximate models of foraging that feeding motivation scales inversely to gut fullness. We plea for a tighter integration between ultimate and proximate foraging models.     

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.     

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。     

Locomotor Ability and Wariness in Yellow-Bellied Marmots

Animals employ a variety of behaviors to reduce or manage predation risk. Often, these are studied in isolation, but selection may act on packages of behavior that are referred to as behavioral syndromes. We focused on yellow‐bellied marmots (Marmota flaviventris) and examined three commonly studied antipredator behaviors. We fitted general linear models to explain variation in maximum running speed, time allocated to vigilance and foraging during bouts of foraging, and flight initiation distance (FID). Marmot maximum running speed was influenced by the substrate run across; marmots ran fastest across dirt or low vegetation and slowest across stones or talus. Incline and several other variables shown to affect running speed in other marmot species failed to explain significant variation in yellow‐bellied marmots. From these results we expected marmots to be sensitive to substrate while foraging, but insensitive to incline. However, time allocated to foraging was affected by incline but not by substrate. In bouts of foraging observed in different habitats, and on different inclines, more time was allocated to foraging and less to vigilance on steep slopes and less on level ground. Substrate influenced FID. Marmots in tall vegetation were less tolerant of an approaching person than were those in shorter vegetation. Finally, we found significant correlations between the residuals from the maximum running speed model and the residuals from the time allocated to vigilance and foraging models. We found a tendency for marmots that ran slower than predicted to be less vigilant while foraging. We also found that relatively slow marmots engaged in more active foraging and less vigilance during foraging bouts. This finding suggests a ‘locomotor ability‐wariness while foraging’ syndrome. It also suggests that vulnerable individuals minimize their exposure while foraging.     

Food choice causes interrupted feeding in the generalist grasshopper Schistocerca americana: further evidence for inefficient decision-making

Individual grasshoppers were observed foraging on either single food plant species or binary mixtures of them. Individuals ate slightly less on the mixtures, but grew faster on them. In the mixtures, individuals fed regularly from both plant species, typically eating both within a single meal. Feeding bouts were longer on the preferred plant species in the mixture. The pattern of foraging behavior was markedly different in mixtures and single food treatments. In mixtures, although the total length of time engaged in ingestion over the day was slightly less than in single food treatments, there were relatively more short feeding bouts and more gaps between feeding bouts within meals, so that the time spent in activities associated with the food was markedly longer on mixtures. This is discussed in terms of efficiency and the problem of choice that faces individual generalists. The efficiency factor and its impact on vigilance is added to the other costs of being a polyphagous herbivore.