Fig Foraging by Dichromatic and Trichromatic <Emphasis Type="Italic">Cebus capucinus</Emphasis> in a Tropical Dry Forest

Figs are important resources for frugivores, and Ficus is an ideal taxon for evaluating patterns of primate foraging related to food color. Ficus spp. can be classified as conspicuous (color change from greenish to reddish during ripening) or cryptic (green throughout ripening). To investigate the effect on foraging of color vision phenotype variation for these 2 types of figs, we conducted a 20-mo study on 4 groups of white-faced capuchins (Cebus capucinus) in the Santa Rosa Sector of the ACG, Costa Rica between May 2004 and September 2008. We genotyped all individuals and collected behavioral data on feeding rates, acceptance indices, and foraging sequences. We found a significant effect of fig type; feeding rates and acceptance indices were higher for conspicuous figs than for cryptic figs, and subjects sniffed cryptic figs more often than conspicuous figs. We also found that dichromats sniffed more figs and had longer foraging sequences than trichromats, especially for cryptic figs. Among 6 subtypes of dichromats and trichromats, monkeys possessing the trichromat phenotype with the most spectrally separated L-M opsin alleles showed the highest acceptance index for conspicuous figs, though there were no differences in feeding rates among phenotypes. We conclude: 1) conspicuous figs are visually salient not only for trichromats but also for dichromats, 2) olfaction is important for evaluating edibility of cryptic figs, and 3) the reliance on olfaction for selecting fruit is greater in dichromats. These results indicate divergent foraging strategies among color vision phenotypes for assessing food items.     

Fruit-feeding behaviour and use of olfactory cues by the fruit bat<Emphasis Type="Italic">Rousettus leschenaulti</Emphasis>: an experimental study

This paper describes the feeding behaviour ofRousettus leschenaulti Desmarest, 1820 on lychees, the preferred cultivated food of this bat in captive conditions. We found that feeding comprised 25–30% of the total activity of these animals in a flight cage and that feeding durations were not significantly different between two sexes. To evaluate the role of odor and vision in foraging behaviour, we provided animals with artificial lychees, real lychees and artificial lychees soaked in the juice of real lychees and we recorded the number of feeding approaches to the different “fruit” types. The results indicated that bats approached real fruit significantly more than artificial fruit, and that the number of approaches to the soaked artificial fruit was also significantly higher than to the unsoaked artificial fruit. There were no significant differences between sexes in approach rates to any “fruit” type. We discuss the role of different sensory cues in the foraging behaviour of these bats and emphasize that the olfactory cue is important in detecting food resources and discriminating between different kinds of food items.     

Differences in Activity Budgets and Diet Between Semiprovisioned and Wild‐Feeding Groups of the Endangered Barbary Macaque (Macaca sylvanus) in the Central High Atlas Mountains,Morocco

The Barbary macaque, Macaca sylvanus is a very adaptable primate species occupying a wide range of habitats in Morocco and Algeria. Several groups of this endangered macaque can be found in tourist sites, where they are affected by the presence of visitors providing food to them. We compare the activity budgets and the diet of semiprovisioned and wild‐feeding groups of Barbary macaques in the central High Atlas Mountains of Morocco from February to August 2008. We used instantaneous scan sampling at 15‐min intervals. The behaviors included in the activity budget were feeding, moving, foraging, resting, and aggressive display. Food items were grouped into seven categories. We found no differences between the two groups in the daily percentages of records attributed to feeding. The semiprovisioned group spent significantly more time engaged in resting and aggressive behavior, and foraged and moved significantly less than the wild‐feeding group. There was no significant difference between the two groups in time spent eating leaves, fruits, or roots and bark. The semiprovisioned group, however, spent significantly less time per day feeding on herbs, seeds, and acorns than the wild‐feeding group. Human food accounted for 26% of the daily feeding records for the semiprovisioned group and 1% for the wild‐feeding group. Our findings agree with previous studies and indicate that in the tourist site, where food is highly clumped, macaques decreased foraging time yet showed higher levels of contest competition. Our results support the common claim that the diet of the Barbary macaque is highly flexible, differing among its varied habitats. Conservation efforts for the Barbary macaques should take into account the changes in behavior that human‐modified environments may cause.     

Foraging behaviour of non-breeding Western Sandpipers Calidris mauri as a function of sex, habitat and flocking

Individuals within a population may vary considerably in the way they exploit available food resources. If the sexes differ in the size of their feeding apparatus, there can be differences in foraging behaviour and habitat use, hence one sex may be more susceptible to competition. We examined relationships between sexual dimorphism in bill size and foraging behaviour, and habitat and microhabitat use of non-breeding Western Sandpipers Calidris mauri at Bahía Santa María, northwestern Mexico. Western Sandpipers are sexually dimorphic, with females about 15% longer-billed than males. Males used a higher proportion of pecks, had a lower probing–pecking rate, walked at higher rates, foraged at sites with lower water content, and had greater variation in foraging technique than females. Moreover, males decreased their proportion of pecks and foraged at a higher rate than females when they changed from feeding alone to feeding in flocks, suggesting a greater safety advantage or susceptibility to conspecific interference when flock feeding. We compared behaviour and microhabitat usage in three habitats: brackish flats, mangroves, and cattail marshes. Sex-related differences in foraging behaviour and microhabitat use were consistent among habitats. Birds in brackish flats and mangroves used a higher proportion of pecks, foraged at lower rates and walked at higher rates, and foraged at deeper sites, with a lower proportion of water cover, than those in cattail marshes. Sex-related differences in foraging behaviour and microhabitat should reduce the level of competition between sexes, and may account for sex differences in Western Sandpiper distribution observed between habitats in Bahía Santa María.     

Intersexual variation in the foraging ecology of sexually monochromatic Western Wood‐Pewees

ABSTRACT Investigators generally pool observations of males and females in studies of the foraging behavior of sexually monochromatic songbirds. However, such pooling can obscure possible intersexual differences. We compared the foraging behavior of male and female Western Wood‐Pewees (Contopus sordidulus), a sexually monochromatic species, in the Sierra Nevada Mountains of California during the breeding seasons of 2007 and 2008. We recorded 143 foraging observations (male N= 74, female N= 69). Overall, mean foraging rates of females (2.8 attacks/min) were higher (P < 0.001) than those of males (1.1 attacks/min). In addition, female foraging rates were significantly higher during incubation than during the nest building, nestling, and fledgling periods. When foraging, males perched higher above ground than females (means = 17.1 and 6.7 m, respectively). Differences between male and female Western Wood‐Pewees in foraging rates and perch heights suggest that males may spend more time on vigilance while females focus on foraging quickly during incubation and when feeding nestlings. Because metrics such as foraging attack rates are sometimes used as indicators of habitat quality and we found that rates can differ between the sexes and among nesting stages, investigators should consider the possibility of such differences when assessing habitat quality, especially for sexually monochromatic species of birds.     

Foraging Behaviour of Subordinate Great Tits (Parus major). Can Morphology Reduce the Cost of Subordination?

This paper studies the magnitude of the behavioural shift, from forage standing to forage hanging, of subordinate great tits ( Parus major ) in two different social contexts: feeding solitarily vs. feeding with a dominant conspecific. The aim is to test the hypothesis that differences in morphological design provide subordinates with varying abilities to reduce the presumed costs of subordination. We find that different subordinate individuals change the foraging behaviour, occupying a different niche when an intra-specific competitor is present. Morphology linked to sexual dimorphism, specifically body mass, is the factor responsible for the different magnitudes of change. Lighter subordinates can remain longer than heavier ones at the feeding patch without interrupting their foraging. Thereby, the former reduce the costs of being subordinate more than the latter. Among subordinates, females are lighter than males; they also spend more time feeding in the presence of a dominant conspecific than males do. No differences are found between age categories. We find no relationship between tarsus length and individual ecological plasticity. Our results support the idea that the ecological plasticity due to morphological differences is a mechanism that allows subordinate individuals to overcome costs associated with subordination.     

实验检验东方田鼠学习能力的性别差异

东方田鼠选择从禾本科植物叶片的不同部位采食会引起食物摄入率的巨大差异,这项具有明显的认知和学习行为的觅食活动是否存在性别差异。在由带两片叶的新鲜马唐茎秆配置的密集食物斑块上,测定东方田鼠觅食马唐叶片的行为序列、过程及参数,检验其在学习能力上的性别差异。结果表明,雌鼠和雄鼠的觅食决定时间尽管均随觅食学习天数的增加而减小,但雌鼠的觅食决定时间显著地高于雄鼠的;雌鼠和雄鼠均由从对叶片部位无差别选择和采食转向偏爱从叶片基部采食。学会这项采食技能后,雌鼠从叶片基部采食的选择指数显著地高于雄鼠的;二者的摄入率均随觅食学习天数的增加极显著地增大,但雌鼠的摄入率却极显著地高于雄鼠的。研究结果充分验证了提出的假设:雄鼠较雌鼠能更快地习得从禾本科植物叶片的基部采食和收获食物;但当二者习得这项采食技艺后,雌鼠较雄鼠能获得更高的食物摄入率;同时也说明,新建立的评价小型植食性哺乳动物认知和学习能力的实验方法,具有可靠的有效性和实用性。     

Temporal and areal feeding behavior of the butterflyfish,Chaetodon trifascialis,at Johnston Atoll

Synopsis The chevron butterflyfish,Chaetodon trifascialis, is found throughout the Indo-Pacific. It is a territorial, diurnal, corallivore found in close association withAcropora spp. corals. The feeding behavior of 33 individuals was studied over six seasons in three habitats.Chaetodon trifascialis spent one third of its active time feeding. However, there was much individual variation. Fish had significantly higher feeding rates during the early afternoon, and there were no significant differences in the feeding rates between the seasons. Feeding rates were significantly different between the three habitats. TheMontipora-rich habitat had the highest feeding rates (x = 10.74 bites min^-1 ± 0.87, all corals combined) and theAcropora-Montipora mixed habitat had the lowest feeding rates (x = 4.58 bites min^-1 ± 0.63, all corals combined). Females fed significantly more than males. WhileC. trifascialis had been thought to only eatAcropora spp. corals, it occasionally fed onMontipora spp. andPocillopora sp. corals whenAcropora spp. were scarce.Chaetodon trifascialis exhibited patterns predicted by foraging theory of an energy maximizer. Territory sizes were inversely related to food density and feeding rates were inversely related to intruder rates. This is a promising system for future testing of foraging strategy models.     

Variation in foraging behavior facilitates resource partitioning in a polymorphic cichlid, Herichthys minckleyi

We examined foraging behavior (microhabitat use and feeding behavior) in a trophically polymorphic cichlid fish, Herichthys minckleyi, to address several questions regarding resource partitioning in this threatened species. These include: (1) do morphotypes demonstrate different foraging behaviors? (2) do individuals within a morphotype vary in their foraging behavior (e.g. are some individuals specialists, only using a subset of available resources, while other are generalists)? (3) do foraging behaviors vary between isolated pools? (4) do foraging behaviors vary across seasons? We quantified microhabitat use and feeding behavior for over 100 individuals (of two morphotypes) feeding freely in two isolated pools (populations) and across two seasons (winter and summer). We found differences in foraging behavior between morphotypes and individual specializations within morphotypes; i.e. some individuals specialize on certain food resources by using a few feeding behaviors within a subset of microhabitats, whereas others employ a range feeding behaviors across many microhabitats. Foraging behavior also varied between pools and across seasons. This spatial and temporal variation in foraging behavior and resource use may serve to maintain this polymorphism, as the relative fitness of the each morph may vary over space and time.     

Foraging and habitat use of common duikers,Sylvicapra grimmia,in a heterogeneous environment within the Soutpansberg,South Africa

We investigated aspects of the foraging behaviour and activity patterns of free‐ranging common duikers (Sylvicapra grimmia) within the Soutpansberg, South Africa. We used giving‐up densities (GUD) and camera traps to test for habitat selection and patch‐use behaviour by common duikers inhabiting a grassland containing distinct ‘islands' of woody vegetation. Foraging in or around a wooded island was affected by its surrounding vegetation. GUDs were significantly lower in portions dominated by tall grass and scattered ferns and highest in areas with open short grass and thick fern. Using grids of 5 × 5 stations, we mapped the duikers' foraging on a larger scale that incorporated neighbouring rocky hillsides. The duikers preferred feeding in areas with tall grass and scattered fern (sufficient cover and escape routes), followed by the wooded islands and thick fern (lack of sightlines/escape routes and presence of predator‐ambush sites), whereas little foraging occurred at the edges and rocky areas (hard substrate that impede escape potential). Photos (total 873) revealed solitary activity, highest in the late afternoon. Photos of vigilant individuals were mostly from rocky and fern habitats. Our results suggest that the duikers allocated their feeding efforts, activity and vigilance patterns to attune to their perceived risk of predation within their heterogeneous environment.     

Climatic determinants of diet and foraging behaviour in baboons

Baboons (Papio spp.) are characterised by a large degree of variation in foraging behaviour and dietary composition. Previous analyses have suggested that much of this can be traced to differences in ecological conditions between sites. The proximate mechanism underlying these relationships is assumed to be mediated via the impact of climatic conditions on food availability, and ultimately the impact that this has on dietary composition. This paper examines these relationships more explicitly. Data from 15 baboon populations were used to assess the relationship between foraging variables and dietary composition. Only feeding time showed significant relationships with dietary composition, with percentage of time spent feeding decreasing with proportion of fruit in the diet, and increasing with the proportion of subterranean items. No relationships were found between diet composition and moving time or day journey length, although significant relationships were found between these variables and group size. The proportions of feeding time spent feeding on fruit, subterranean items and leaves were functions of the ecological conditions experienced by that population, although no relationships were found for the percentage of feeding time devoted to flowers or animal material. The relationships between the proportion feeding time spent on fruits, leaves and subterranean items and ecological variables could be best explained through understanding the way in which bush and tree level vegetation respond to the climatic environment and the impact this has on fruit availability. In this respect, temperature and seasonality are the key climatic parameters. This provides good support for the idea that the proximate mechanism underlying the relationship between foraging time and ecological variables is mediated via the impact of the climatic environment on vegetation structure and food availability. Similar factors have been proposed to explain much of the geographic variation in species diversity, suggesting that these relationships have far wider relevance and may account for much of the observed geographical variation in mammalian behaviour.     

Differential survival in adult Eurasian oystercatchers Haematopus ostralegus

I explored the fitness implications of individual and sex differences in foraging strategy in the Eurasian oystercatcher Haematopus ostralegus by monitoring the survival of individually colour-ringed birds of known sex and known feeding specialisation. Over the period of this study, adult female annual and overwinter survival was significantly lower than adult male survival. However, contrary to previous findings, no differences in survival were found between birds of different feeding specialisations. Lower female survival was not due to sex differences in feeding specialisation. Possible mechanisms for sex differences in survival and the survival implications of different feeding specialisations are discussed. I conclude that sex differences in survival may be due to differences in social status. I also suggest that worm/clam feeders and mussel-stabbers, feeding specialisations previously associated with lower survival rates, may have benefited more than mussel-hammerers from milder winter temperatures in recent years.     

Sex‐specific food provisioning in a monomorphic seabird,the common guillemot Uria aalge: nest defence,foraging efficiency or parental effort?

Sexual differences in food provisioning rates of monomorphic seabirds are well known but poorly understood. Here, we address three hypotheses that attempt to explain female-biased food provisioning in common guillemots Uria aalge : (1) males spend more time in nest defence, (2) females have greater foraging efficiency, and (3) males allocate a greater proportion of foraging effort to self-maintenance. We found that males spent no more time with chicks than females but made longer trips and travelled further from the colony. There was extensive overlap between sexes in core foraging areas, indicating that females were not excluding males from feeding opportunities close to the colony. However, as a result of their longer trips, the total foraging areas of males were much greater than those of females. There was no difference between sexes in overall dive rate per hour at sea, in behaviour during individual dives or in a number of other measures of foraging efficiency including the frequency, depth and duration of dives and the dive: pause ratio during the final dive bout of each trip, which was presumably used by both sexes to obtain prey for the chick. These data strongly suggest that sexes did not differ in their ability to locate and capture prey. Yet males made almost twice as many dives per trip as females, suggesting that males made more dives than females for their own benefit. These results support the hypothesis that female-biased food provisioning arose from a difference between sexes in the allocation of foraging effort between parents and offspring, in anticipation of a prolonged period of male-only post-fledging care of the chick, and not from differences in foraging efficiency or time spent in nest defence.     

Sex-specific chick provisioning and diving behaviour in the wedge-tailed shearwater Puffinus pacificus

Mechanisms that drive sex-specific foraging behaviour in seabirds are not fully understood. In some cases, sexual-size dimorphism has been implicated. However, recent empirical work indicates that foraging behaviour may also differ between sexes of monomorphic seabird taxa. We simultaneously examined sex-specific differences in adult foraging behaviour, chick provisioning rates and maximum dive-depths in a monomorphic seabird, the wedge-tailed shearwater Puffinus pacificus . We found significant divergence between sexes. Mean foraging trip length was longer, provisioning rate lower and mean maximum dive-depth shallower in females. We found no evidence of divergence in foraging behaviour due to condition-dependant increases in self-provisioning by females, or differences in the nest attendance patterns of each sex. In addition, chick body condition did not influence meal mass or trip length differently in one or other sex. Consistent with results obtained for dimorphic species we suggest that inter-sexual competition at the foraging grounds provides the most parsimonious explanation for the sex-specific differences observed in this monomorphic species. Based on our findings we believe this possibility warrants further critical investigation.     

The foraging ecology of two sympatric gobiid fishes: importance of behavior in prey type selection

Synopsis The foraging ecology of two temperate marine gobies (Pisces: Gobiidae) was studied in rocky subtidal habitats off Santa Catalina Island, California. The bluebanded goby, Lythrypnus dalli, foraged from exposed ledges and fed on planktonic and benthic prey, although planktonic prey were more important in diets by number and weight. The more cryptic zebra goby, Lythrypnus zebra, remained hidden under rocks and in crevices feeding on benthic prey almost exclusively. The active selection of particular prey taxa from the two prey sources (water column and substratum), mediated by species-specific differences in foraging behavior, resulted in interspecific differences in type, number, size and weight of prey consumed. Interspecific differences in foraging ecology reflect the selection of prey most readily available to these fishes that occupy specific and fixed microhabitats within rocky reefs.     

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.     

Effects of light level and growth history on attack distances of visually foraging juvenile salmon in experimental tanks

The effects of light level, developmental pathway, and previous growth history on the foraging attack distances of juvenile Atlantic salmon Salmo salar were examined in circular rearing tanks. Former manipulation of growth rates had no significant influence on distances moved to intercept food items despite the fact that it caused substantial differences in post-treatment growth. Attack distances of fish that were entering a state of overwinter dormancy (lower modal group; LMG) were shorter than those of actively feeding (and growing) fish (upper modal group; UMG). These differences were explained generally by differences in body size between the two groups, suggesting that actual effort per attack was unassociated with growth requirements. Significant differences between growing and non-growing fish in attack distances could contribute to the variation in growth rates through their effect on feeding rates, but were unlikely to have affected energetic costs. This may be due to the fact that attack distances were consistently short throughout the study period as is also evident from the pattern of change between night and daytime. Whereas in the first experiment (daylight v . twilight) fish moved further to reach food during the day, in the second (daylight v . overcast night) nocturnal attack distances matched (LMG fish) or exceeded (UMG fish) diurnal attack distances. Thus diurnal attack distances were probably minimized in the second experiment. These results are interpreted within a framework of overwintering strategies.     

Foraging flight distances as a measure of parental effort in blue tits Parus caeruleus differ with environmental conditions

Life-history theory predicts that parental effort in nestling provisioning is optimised in relation to the quality of individuals and/or their habitat. We studied the investment of breeding pairs of blue tits Parus caeruleus for their reproduction during three breeding seasons in deciduous (high quality) vs. mixed (low quality) habitats in order to quantify to what extent habitat quality affects parental effort. Parental effort (costs) was related to their feeding rates and flight distances during foraging. In the deciduous habitat flight distances between nest and foraging patch were shorter than in the mixed habitat (22 m and 40 m, respectively), but the feeding rates did not differ between the habitats. The total flight distance per breeding pair from the first day after hatching until the 17th day of the nestling period was about half of the distance observed in the mixed habitat (375 km and 674 km, respectively). As the quality of fledglings did not differ between habitats, the higher number of fledglings per brood reflects better rewards per foraging trip in the deciduous than in the mixed habitat. Considering the parental foraging effort (costs) and, the quality and number of offspring (benefits), the benefit-cost-ratio was 2–3 times higher in the deciduous than in the mixed woodland.     

Responses by Central‐Place Foragers to Manipulations of Brood Size: Parent Flickers Respond to Proximate Cues but do not Increase Work Rate

Manipulations of brood size measure the willingness or ability of parents to invest in offspring and different reproductive roles may lead to differences in feeding effort between the sexes. Parental investment in birds is usually assessed by quantifying feeding rates, but this provides an incomplete picture of parental effort because it fails to account for how parents collect food on the landscape. We studied northern flickers (Colaptes auratus), a woodpecker in which males provide the majority of parental care and used a repeated measures design and short‐term (24 h) brood enlargements (N = 35) and reductions (N = 27) to assess effects of treatment on feeding rates to nestlings and parental foraging behaviour. Parents of enlarged broods did not significantly increase feeding rate, resulting in a decline in nestling mass. Parents of reduced broods decreased their feeding rates by 84%, but increased per capita feeding rates, resulting in nestling mass gain. The variation in feeding rates to enlarged broods was not influenced by feather corticosterone, body condition, feather re‐growth rate or mass change between the incubation and nestling periods. Foraging pattern on the landscape remained the same during the enlarged treatment for both sexes. We conclude that flickers respond to proximate cues in brood demands, but do not increase feeding rates to enlarged broods, at least in the short term. A literature review suggested that this lack of response is atypical for short‐lived species. We hypothesize that parents in species with large home ranges and long nestling periods face energetic limitations that constrain their ability to respond to enlarged broods. We encourage future studies to assess foraging behaviour on the landscape to document important trade‐offs for parents such as predation risk and energy expenditure while feeding offspring.     

Flexibility in the Foraging Behavior of Blue Tits in Response to Short‐Term Manipulations of Brood Size

Previous work suggests that short‐term changes in feeding rate are usually produced by the parent‐offspring interaction. However, few studies have properly tested this assumption. In this study, we attempt to explore the short‐term consequences of daily (within‐pair) brood size manipulations (reduced, original, and enlarged) on feeding behavior (provisioning rates, prey size, and prey type) of Mediterranean blue tits Cyanistes caeruleus. Total provisioning rates were lowest when broods were reduced in size and greatest when broods were enlarged. Mean prey size was also affected by the brood size changes: parents tended to bring larger prey when confronted with low brood demand reinforcing the view that a trade‐off exists between minimizing foraging time and maximizing food quantity. Such differences in feeding frequencies and the load sizes delivered may be explained by changes in the parents’ foraging tactic. Increase of brood size compelled parents to work harder and be less selective in prey choice; we found that stressed birds with a high level of feeding responsibility (hungry nestlings) opted to concentrate on more readily available food items (Tortricids). On the other hand, their immediate reaction when faced with a low level of feeding responsibility was to decrease this prey type in the diet, so that the percentage of other preys (Noctuids) in the diet increased. There was no intersexual difference in the way in which parents responded to the manipulation. In sum, our results revealed a flexibility in foraging strategies of blue tits to cope with changing scenarios, which supports the idea that provisioning behavior is largely governed by nestling demand.