Resource base,parity, and reproductive condition affect females' feeding time and nutrient intake within and between groups of a baboon population

Summary We examined within- and between-group differences in aspects of feeding and nutrient intake among adult females of a single population of baboons (Papio cynocephalus) in Amboseli National Park, Kenya. Differences in time spent feeding, daily energy and protein intake and feeding efficiency (nutrient intake per minute spent feeding) reflected differences in resource base, reproductive condition and parity. Baboons that partially fed from a lodge garbage dump spent less than half the time feeding than those that were feeding totally in the wild. During this greatly reduced feeding time, the garbage-feeding group had a similar daily energy intake and only a slightly lower daily protein intake relative to wild-feeding baboons. Consequently, the feeding efficiency of the semi-provisioned baboons was appreciably higher than that of the non-provisioned baboons. For the totally wild-feeding baboons, samples were large enough to permit analyses of feeding time and nutrient intake during different reproductive states and parity. Females spent more time feeding and had higher daily energy and protein intake when they were pregnant or lactating than when they were sexually cycling. Nulliparous females spent more time feeding than their multiparous counterparts. The daily energy intake of nulliparous females was higher than that of their multiparous counterparts, but their daily protein intakes did not differ significantly. Pregnant or lactating and nulliparous females had higher feeding efficiency than their sexually cycling and multiparous counterparts. The two nulliparous females in the garbage-feeding group spent more time feeding but did not take in more energy or protein per day than their multiparous counterparts.     

Prey recognition time of praying mantids (Dictyoptera: Mantidae) and consequent survivorship of unpalatable prey (Hemiptera: Lygaeidae)

When juvenile praying mantids (Tenodera sinensis)were exposed to unpalatable prey (the milkweed bug Oncopeltus fasciatus),they attacked, sampled, and then usually rejected the prey. About 70% of the handling time was spent feeding. When offered a second milkweed bug, the mantids usually attacked the prey. However, the overall time required for the mantids to sample, recognize, and then reject the unpalatable prey decreased by half. The proportion of handling time that was spent feeding remained the same as in the first encounter. In contrast, when the second prey individuals encountered by mantids were Drosophila melanogaster,the flies were completely consumed and the proportion of handling time that was spent feeding significantly increased. When praying mantids were exposed to the milkweed bugs for the first time, up to 33% of the bugs survived attack by the mantids. Survival of milkweed bugs increased to 55% when mantids had been previously exposed to the bugs. In contrast, flies that were caught never survived.     

Allocating time between feeding, resting and moving by the two-spotted spider mite, Tetranychus urticae and its predator Phytoseiulus persimilis

This study characterizes the timing of feeding, moving and resting for the two-spotted spider mite, Tetranychus urticae Koch and a phytoseiid predator, Phytoseiulus persimilis Athias-Henriot. Feeding is the interaction between T. urticae and plants, and between P. persimilis and T. urticae. Movement plays a key role in locating new food resources. Both activities are closely related to survival and reproduction. We measured the time allocated to these behaviours at four ages of the spider mite (juveniles, adult females immediately after moult and adult females 1 and 3 days after moult) and two ages of the predatory mite (juveniles and adult females). We also examined the effect of previous spider mite-inflicted leaf damage on the spider mite behaviour. Juveniles of both the spider mite and the predatory mite moved around less than their adult counterparts. Newly emerged adult female spider mites spent most of their time moving, stopping only to feed. This represents the teneral phase, during which adult female spider mites are most likely to disperse. With the exception of this age group, spider mites moved more and fed less on previously damaged than on clean leaves. Because of this, the spider mite behaviour was initially more variable on damaged leaves. Phytoseiulus persimilis rested at all stages for a much larger percentage of the time and spent less time feeding than did T. urticae; the predators invariably rested in close proximity to the prey. Compared to adult predators, juveniles spent approximately four times as long handling a prey egg. The predator-prey interaction is dependent upon the local movement of both the predators and prey. These details of individual behaviours in a multispecies environment can provide an understanding of population dynamics.     

Prey feeding increases water stress in the omnivorous predator Dicyphus hesperus

The effects of water stress (produced by water deprivation and prey feeding) on plant feeding were investigated in the omnivorous predator Dicyphus hesperus Knight (Hemiptera: Miridae). The objective was to determine if prey feeding aggravated water deficits and thus increased plant feeding. We measured plant feeding in a factorial experiment where female D. hesperus were prepared for experiments by providing or withholding water and/or prey for 24 h. We then evaluated the amount of plant feeding on Nicotiana tabacum seedlings by the direct observation of insects at three different densities of the prey, Ephestia kuehniella eggs. The amount of plant feeding, as measured by frequency of plant feeding bouts and time spent plant feeding during observation, was significantly greater for water‐deprived individuals than for those that had been provided with water. Individuals that had been provided with prey fed on plants at a significantly higher frequency than prey‐deprived individuals at two of the prey densities used in the experiment. These results support the hypothesis that plant feeding in zoophytophagous Hemiptera facilitates prey feeding by providing water that is essential for predation.     

Foraging in yellow dung flies: testing for a small-male time budget advantage

1. Mating and foraging are generally mutually exclusive activities. Individuals are thus faced with a continuous trade-off between time and energy expended in foraging and mating, but different phenotypes should respond to this trade-off in different ways. 2. Sexual selection theory predicts that females should maximize their time and energy spent gathering resources, whereas males should maximize their time and energy spent obtaining mates, thus minimizing their time spent foraging, subject to the constraint that they need to forage minimally to sustain their activity. 3. Smaller individuals require less food to maintain their activity. Small males in particular could therefore increase mating effort at the expense of foraging effort and, all else being equal, may thus enjoy a time budget advantage relative to large males. On the other hand, larger individuals may compensate by being more efficient at finding prey and/or extracting nutrients. 4. The effects of sex, body size, and prey density on foraging time budgets of male and female yellow dung flies, Scathophaga stercoraria, were investigated in the laboratory. 5. Higher prey density (Drosophila melanogaster) resulted in reduced feeding (= handling) and hunting (= waiting or search) times for both sexes, as predicted by the marginal value theorem applied to foraging theory. Females fed longer on a prey item than did males, and also caught the next prey item more quickly. Large individuals extracted nutrients more quickly, but were not faster at catching prey. Small individuals satiated more quickly than larger individuals and also ate fewer prey items. 6. These results are largely consistent with the predictions and suggest a small-male time budget advantage in the yellow dung fly. Integrating the various predictions to test directly for a small-male time budget advantage is difficult in the laboratory, however, because hunting times are unlikely to reflect the natural situation. To what extent these results lead to increased probabilities for small males of obtaining matings in the field remains to be demonstrated.     

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.     

Quantitative field investigations of feeding and territorial behaviour of young-of-the-year brook charr,Salvelinus fontinalis

Synopsis Direct observations of young-of-the-year brook charr, Salvelinus fontinalis, in a second-order woodland stream indicated that most of their feeding effort was directed toward sub-surface, drifting prey (83% of feeding time). Feeding from the substrate and water surface was much less frequent (17% of feeding time). Comparisons of gut contents to drift net and substrate fauna samples corroborated that the most commonly consumed prey (chironomid and trichopteran larvae, ostracods, and ephemeropteran nymphs) were captured primarily from sub-surface, invertebrate drift. The disproportionate numbers of some prey species in the guts of several fish indicate that some prey selection occurred. Territories appeared to be cardioid-shaped, and were often contiguous, with dominance hierarchies evident among the residents. Agonistic interactions were frequent. Charges and chases predominated (91% of interactions) while lateral displays were infrequent (9% of interactions). Overall, these fish spent most of the daylight hours station-holding (77%) and feeding (18%). While only 3% of total time was spent in aggression, this amounted to 14% of the time a fish spent away from its station. There was some indication that territories were defended at a cost of feeding time.     

Changes in the activity budget of cycling female chimpanzees

This study is a preliminary report on the time allocated to various activities by female wild chimpanzees (Pan troglodytes schweinfurthii) during their sexual cycle. Cycling females with maximal tumescence (estrous females) tended to spend more time moving than cycling females with quiescent sexual skin (anestrous females). Although there was no statistically significant decrease in any specific activity that corresponded to the increase in time spent moving, feeding time did decrease in four of the five females. The frequency of approach by females toward males and the frequency of approach by males toward females significantly increased when females were in estrus. Direct aggression by males occurred more frequently toward estrous females than toward anestrous females. The copulation frequency and the frequency of approach to males was not significantly correlated with the increase in time spent moving. There was a high but not significant correlation between the time spent moving and the frequency of direct aggression by males toward females. Mating effort, feeding competition, male aggression, and other possible reasons that might explain the increase in moving time are discussed. Am. J. Primatol. 46:157–166, 1998. © 1998 Wiley-Liss, Inc.     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Estimation of food intake and ingested energy in Daubenton’s bats (Myotis daubentonii) during pregnancy and spermatogenesis

We studied food intake of and estimated ingested energy in female and male Myotis daubentonii during the periods of pregnancy (period 1, 8 May–4 June) and of intense spermatogenetic activity (period 2, 24 July–22 August) over 8 years (1996–2003) in central Germany. We used radiotelemetry to determine the time spent foraging and marked animals with chemiluminescent light-sticks to determine prey attack rates. Body length, body mass, moisture content, and caloric content of chironomids, the main prey of Daubenton’s bats, were measured to estimate the nightly food intake and, in consequence, energy intake. Pregnant females spent significantly more time foraging than males during period 1 and females during the post-lactation period. In contrast, male foraged longer during the period of highest spermatogenetic activity than during late spring and also significantly longer than post-lactating females. Based on a mean number of 8.3 prey attacks per minute, the time spent foraging, and a capture success rate of either 50 or 92%, calculated intake values with a feeding rate of 7.6 insects per minute (=92% capture success) were more consistent with literature data for other insectivorous bats than that of values calculated on the basis of a capture success rate of 50%. In the high capture-success model, calculated insect intake of female bats was 8.0 g during pregnancy and 4.9 g per day during post-lactation, providing 5.0 and 3.0 kJ of ingested energy per gram body mass per day. Calculated intake of male bats was 3.6 g insects per day during late spring and 8.0 g during period of intensive spermatogenesis, providing 2.6 and 5.7 kJ of ingested energy per gram body mass.     

Behavior, Diet, and Movements of the Sulawesi Crested Black Macaque (Macaca nigra)

We present the first field study of activity budgets, diet and ranging patterns of the Sulawesi crested black macaque, Macaca nigra, one of seven macaque species endemic to Sulawesi, Indonesia. We studied three crested macaque groups, ranging in size from 50 to 97 individuals, for 18 months in the Tangkoko-DuaSudara Nature Reserve, North Sulawesi. They spent 59% of the day moving and procuring food, especially fruits, and 41% of the day resting and socializing. Their diet is composed of more than 145 species of fruit (66% of observed feeding bouts), vegetative material (2.5%), invertebrates (31.5%), and occasional vertebrate prey. Group differences were more pronounced than seasonal or diurnal differences. Specifically, the largest group moved farther during the day, moved at a faster and more uniform rate, ate less fruit, rested more, and socialized less than the smaller groups did. The largest group had the largest home range, but it included less primary forest and more disturbed habitat than the ranges of smaller groups. There are individual differences in activity budgets of adult males and females in time spent moving, resting, feeding, and socializing that may reflect differences in reproductive strategies of males versus females. The behavior of large juveniles is more similar to that of adults than to that of small juveniles. Daily movements and use of home range are correlated with diet. Macaques moved shorter distances as the proportion of time spent feeding on fruit increased, and the top four dietary items accounted for most of the variance in entry into hectare blocks of home range.     

Influence of Human Food Consumption on the Time Budget of Vervets

In nonhuman primate groups that utilize crop and food-raiding strategies, the inclusion of human food in the diet can have a pervasive impact on the activity budget. Human food is usually of higher quality and energy per unit than wild food. Thus, metabolic demands may be reached sooner on a human food diet, which in turn leads to improved foraging efficiency. In particular, a reduction in time spent feeding is associated with an increase in time spent resting. We investigated changes in time budget in a group of vervets (Chlorocebus aethiops pygerthrus) living in a tourist and cultivated area of Entebbe, Uganda (Zoo group). Saj collected 123 hr of focal observations on 17 individuals. Human food constituted half of the diet of Zoo group. Access to human food resulted in a high proportion of time resting and a low proportion of time feeding. The group had a small average daily range and a small home range in relation to other reports on unprovisioned groups. The time budget differences suggest that the juveniles spent more time moving and feeding and less time resting than adults did, and that adult females were more social than adult males were. These patterns have been observed in both provisioned and unprovisioned vervet groups. In contrast to a report from an unprovisioned group (Harrison, 1983), lactating females in this provisioned situation were able to maintain feeding and activity levels similar to those of other adults. However, adult females spent significantly less time consuming human food than juveniles did. We suggest that the risk associated with human food consumption may explain the low consumption of human food by females with dependent offspring. The differences may also reflect the different nutritional requirements of lactating females or the tendency for juveniles to be more exploratory than other group members.     

Age- and dominance-related variation in feeding time among free-ranging female rhesus monkeys

Observations were made on the feeding behavior of 34 lactating rhesus females (Macaca mulatta) at Tughlaqabad, India. Consistent interindividual differences in feeding times were found and two sources of this variation were identified. Time spent feeding was found to decline with female age; rhesus mothers 4–6 years old spent over 40% more time feeding than did females 11–15 years of age. When the animals were divided into high- and low-status groups, dominance was also found to relate to individual feeding times; high-status females spent more time feeding that did low-status females, but this difference was evident only when the effect of female age was held constant. The age effect is probably an expression of age-related variation in nutritional requirements due to growth in the youngest subject and, perhaps, a declining basal metabolic rate in the oldest. We speculate that the comparatively weak dominance effect is due less to the aggressive interruption of feeding bouts than it is to a presumed greater vigilance on the part of subordinant mothers in monitoring their infants' whereabouts.     

Activities and feeding behavior of free-ranging pregnant baboons

The activities and feeding behavior of pregnant baboons, Papio cynocephalus,were studied in two free-ranging groups in Amboseli National Park, Kenya. Both the environmental conditions and the age of their fetuses influenced females’ activity budgets and feeding behavior. Females fed more and rested less during the dry season than during the wet season, and most females spent progressively more time feeding and less time grooming than expected (based upon the amount of recent rainfall) as their fetuses matured. During the wet season, females also devoted significantly greater proportions of feeding time to consumption of grass blades, leaves, and flowers and significantly smaller proportions of their feeding time to feeding upon seeds, corms, and fruit. As their fetuses became older, the proportion of time spent feeding upon seeds increased, and the proportion of time spent feeding upon grass blades and fruit declined.     

Effect of delayed mating and prolonged engorgement on the reproductive fitness of female Amblyomma limbatum (Acari: Ixodidae) in marginal population areas

Reptile ticks mate while females are attached to their host. Following mating, females engorge, detach and then lay their eggs. This study examines whether the time that females of the reptile tick Amblyomma limbatum spend on hosts prior to mating and the time they take to engorge on hosts after mating influence their reproductive fitness, as measured by the number of viable eggs they produce. When compared with females that experienced no temperature-induced delay in mating, females attached for an initial period to hosts with body temperatures too low to induce mating had no decline in their reproductive fitness, once provided with suitable temperatures for mating. However, on hosts with suitable body temperatures for mating, female ticks that took longer to mate, took longer to engorge and had reduced reproductive fitness. In addition, females that spent more than 20 days feeding on hosts after mating had reduced reproductive fitness, irrespective of the time they spent on hosts prior to mating compared with females with shorter feeding periods. Thus, the time that A. limbatum females spent on hosts prior to mating and/or after mating has a significant bearing on the number viable progeny produced. The ecological implications of these results are discussed in relation to females colonizing marginal population areas at the edge of their distributional range.     

Prey preference and consumption capacity of Nephus arcuatus (Coleoptera: Coccinellidae): the influence of prey stage,prey size and feeding experience

This study examined the feeding habits of Nephus arcuatus Kapur (Coleoptera: Coccinellidae), an important predator of mealybugs in south-western Iran. The consumption capacity of male and female N. arcuatus adults was determined by their feeding on eggs, first-instar nymphs, and adult females of two destructive mealybugs, Nipaecoccus viridis (Newstead) and Planococcus citri Risso, over a 24-h period. N arcuatus consumed significantly more first-instar nymphs than eggs, and more eggs than female adults of both prey species. In addition, we also studied the developmental stage prey preference of adults reared on either N. viridis or P. citri and found that the prey preference of females did not change with the developmental stage of the mealybug. Meanwhile, the males reared on either N. viridis or P. citri showed a significant preference for the first-instar nymphs of P. citri over first-instar nymphs of N. viridis, while a preference for the eggs or adult females of these two mealybugs was not observed. This selection of first-instar nymphs by males was not tied to its previous feeding experience. Our findings suggest that prey stage, prey size and previous feeding experience had no effect on the prey selection of N. arcuatus, making it a good candidate for the biological control of mealybugs.     

Parental care behavior in the monogamous,sexually dimorphic Madagascar paradise flycatcher: sex differences and the effect of brood size

I studied the parental care behavior of the Madagascar paradise flycatcher Terpsiphone mutata in northwestern Madagascar. I especially focused on feeding, brooding and vigilance behaviors. Feeding rate did not differ between males and females, but females spent more time at the nest than males. Females dedicated their time to brooding, while males perched on the nest and were vigilant. Both parents changed the feeding rate in relation to brood size, so the feeding rate per nestling was not different among nests of different brood size. Duration of brooding by females increased with decreasing brood size, suggesting that the Royama effect, the pattern of lower feeding rate per nestling in larger broods, did not apply in this study. Males spent more time on vigilance than females. Anti-predator vigilance by males should be important for nestling survival given the high predation pressure typical of this population. In conclusion, males provide considerable parental care probably to minimize nestling starvation and to avoid nest predation. My results are not consistent with the general pattern of less parental effort by males in monogamous, sexually dimorphic species.     

Feeding behavior ofEuseius stipulatus andTyphlodromus phialatus on the citrus red mitePanonychus citri [Acari: Phytoseiidae,Tetranychidae]

Observations were made on the feeding behavior of the two main phytoseiid species in Spanish Citrus orchards,Euseius stipulatus (Athias-Henriot) andTyphlodromus phialatus Athias-henriot. The experiences were carried out by rearing the predatory mites on excised orange leaves, and always with an excess of the prey the Citrus Red Mite (=CRM)Panonychus citri (McGregor). In experiments with all stages of CRM, the number of prey killed per hour was 5.12 and 2.00, the percentage of successful attacks, 58% and 21%, and the mean time spent feeding on each prey was 5.1 and 12.2 minutes for starving females ofE. stipulatus andT. phialatus respectively.E. stipulatus feeds on all stages of the prey except eggs, andT. philatus, on all stages, except males. Both species attack much less successfully females ofP. citri rather than immatures. In experiments with adult females and eggs ofP. citri as prey, the mean number of prey killed daily was 4.51 females forE. stipulatus, and 2.01 females and 2.12 eggs forT. philatus. Considering this killing rate and the number of eggs laid by the predators in the same period, it can be concluded thatE. stipulatus consumes only 30% of the content of the preys killed, whereasT. philatus consumes a percentage of prey variable between individuals and ranging from 40% to 100%. These differences in feeding behavior between the two species could partly explain differences in their efficiency as biocontrol agents ofP. citri observed in the field.      

Food fight: sexual conflict over free amino acids in the nuptial gifts of male decorated crickets

In decorated crickets, Gryllodes sigillatus, the spermatophore that a male transfers at mating includes a gelatinous spermatophylax that the female consumes, delaying her removal of the sperm‐filled ampulla. Male fertilization success increases with the length of time females spend feeding on the spermatophylax, while females may benefit by prematurely discarding the spermatophylaxes of undesirable males. This sexual conflict should favour males that produce increasingly appealing spermatophylaxes, and females that resist this manipulation. To determine the genetic basis of female spermatophylax feeding behaviour, we fed spermatophylaxes to females of nine inbred lines and found that female genotype had a major influence on spermatophylax feeding duration. The amino acid composition of the spermatophylax was also significantly heritable. There was a positive genetic correlation between spermatophylax feeding duration and the gustatory appeal of the spermatophylax. This correlation suggests that genes expressed in males that produce more manipulative spermatophylaxes are positively linked to genes expressed in females that make them more vulnerable to manipulation. Outbred females spent less time feeding on spermatophylaxes than inbred females, and thus showed greater resistance to male manipulation. Further, in a nonspermatophylax producing cricket (Acheta domesticus), females were significantly more prone to feeding on spermatophylaxes than outbred female Gryllodes. Collectively, these results suggest a history of sexually antagonistic coevolution over the consumption of nuptial food gifts.     

烟粉虱天敌沙巴拟刀角瓢虫与越南斧瓢虫的捕食行为比较

在实验室条件下研究了沙巴拟刀角瓢虫Serangiella sababensis Sasaji与越南斧瓢虫Axinoscymnus apioides Kuzne tsov and Ren对烟粉虱Bemisia tabaci(Genna dius)卵的捕食行为及各种行为的时间分配。2种瓢虫的捕食过程非常相似,大致可以分为搜寻、取食、清洁、静息、整翅和排泄6个部分。在整个捕食过程中,沙巴拟刀角瓢虫在饥饿状态下,其各行为所占的时间为爬行时间>取食时间>静息时间>清洁时间>猎物处置时间,而在非饥饿状态下,各行为所占的时间为爬行时间>静息时间>取食时间>清洁时间>猎物处置时间;对于越南斧瓢虫来讲,饥饿状态下各行为所占的时间比例为爬行时间>静息时间>取食时间>清洁时间>猎物处置时间,而在非饥饿状态,越南斧瓢虫用在取食方面的时间很少,仅长于处置猎物的时间。2种瓢虫相比,沙巴拟刀角瓢虫用在爬行和取食方面的时间要比越南斧瓢虫长,而用在静息和清洁方面的时间比越南斧瓢虫短。