Self-selection and utilization efficiency of Heliothis zea larvae feeding on seeds and the walls of snap bean pods

First instar Heliothis zea larvae tended to feed on the leaves of snap beans, but later instars preferred to burrow in the pods. Fifth instars offered only leaves grew poorly because they ate them in small quantities, presumably because of deterrency. The utilization efficiency and growth of fifth instars fed pulp was slightly but significantly inferior to that of seed-fed larvae but was greatly superior to that of leaf-fed larvae. Since the dry matter content of pulp is less than one-third that of seeds, larvae allowed to eat only pulp had abnormally large fresh weight intakes and devoted about three times as much time to feeding as did seed-fed larvae. Larvae offered a mix of seeds and pulp ate both and had a total fresh weight food intake somewhat greater than that of seed-fed larvae but much less than that of pulp-fed larvae. Eating both seeds and pulp offers no obvious nutritional advantage but presumably offers a significant ecological advantage. It is argued that the number of foraging trip a larva must make to discover pods is minimized by feeding on a mix, thus reducing exposure to mortality factors such as predation.     

Foraging and provisioning in Antarctic fur seals: interannual variability in time-energy budgets

This study examined three competing hypotheses to explain howlactating Antarctic fur seals (Arctocephalus gazella) respondto changes in the level of resource availability. Antarcticfur seals have episodic bouts of suckling (1-3 days), alternatingwith foraging trips (3-10 days). Foraging time budgets variedsignificantly (p <.001) among 8 consecutive years at BirdIsland, South Georgia. Foraging trip duration increased during periodsof relative food shortage. Time spent ashore was more consistentamong years than foraging trip duration but declined duringa year of particularly low food availability. In 4 of the 8years, there was a significant positive correlation betweentime spent ashore and foraging trip duration. In the other years,the relationship was close to statistical significance. Energydelivery to pups during suckling bouts followed an asymptoticpower function. Energy gain during foraging trips was estimatedfrom diving behavior, which suggested that the energy gain functionwas linear. Distance traveled during foraging trips was correlatedwith foraging trip duration, and long foraging trips were associatedwith reduced foraging intensity. There was support for the hypothesisthat lactating Antarctic fur seals compensate for reduced resources byincreasing the foraging trip duration rather than working harderand increasing their energy expenditure. However, there wasmost support for the hypothesis that lactating Antarctic furseals adjust time spent ashore as well as foraging trip duration,possibly to maximize the delivery of food to their offspring.Lactation appears to impose constraints on provisioning of offspringthat differ from those of seabirds foraging in the same environment andoften on the same prey.     

A fine-scale time budget of Cape gannets provides insights into the foraging strategies of coastal seabirds

Central-place foragers organize their feeding trips both to feed themselves and to provide their offspring with food. In seabirds, several long-range foragers have been shown to alternate long and short trips to balance these dual needs. However, the strategies of short-range foragers remain poorly understood. We used a precise, miniaturized motion sensor to examine the time budget of 20 breeding Cape gannets, Morus capensis, foraging off the coast of South Africa. Birds stayed at sea for 5.5-25.3 h, occasionally spending the night at sea. The large number of isolated dives and extended flight time observed during these overnight trips suggested that birds either experienced poor foraging conditions or exploited more distant, yet more profitable prey patches. Conversely, birds that stayed at sea for less than 1 day had relatively consistent activity patterns. Most of these birds (88%) foraged actively at the beginning and at the end of the foraging trip. These feeding bouts were separated by protracted periods of sitting on the sea surface. Such resting periods probably allow birds to digest the food ingested during the first part of the foraging trip, so they initially feed themselves, and then obtain food for their chick on the way back to the breeding site.     

Foraging by <Emphasis Type="Italic">Drosophila melanogaster</Emphasis> Larvae in a Patchy Environment

The food resources of Drosophila comprise decaying vegetable matter distributed in patches, yet foraging behavior has not been examined in larvae reared continuously in a patchy environment. Here, the extent and rate of inter-patch movement was studied in larvae of four wild strains of D. melanogaster inhabiting an experimental arena from the egg stage to the third larval instar. The hypotheses were that larvae would forage primarily in the third instar, that larvae would move from low-protein patches at higher rates than from high-protein patches, and that foraging rates would be higher on an agar substrate than on sand. Larvae hatching on a nutrient-poor food patch switched to a nutrient-rich patch during the first instar. The rate of interpatch switching increased with larval age, as did the number of larvae roving on the substrate between food patches. Inter-patch distance affected switching speed---the closer the patches, the faster the switching. High protein patches were preferred over low-protein patches, but there was a bias towards staying on the natal patch. Significant variation among strains in latency to forage, in proportion of larvae that switched patches, and in the rate of roving between patches suggests that there is natural genetic variation for these traits. Larvae switched food patches on a substrate of moist sand as quickly as on an agar substrate.     

Daily activity and minimum food requirements during winter for gentoo penguins (Pygoscelis papua) in the South Shetland Islands, Antarctica

Estimates of daily activity and consequent demand for food during winter are scarce for many polar seabirds, yet essential for assessing constraints on foraging effort, demand for food, and potential competition with local fisheries. We affixed archival temperature tags to gentoo penguins (Pygoscelis papua) from two colonies in the South Shetland Islands to measure the frequency, timing, and duration of foraging trips and to estimate minimum food requirements during winter. Foraging trip frequencies ranged from 0.85 to 1.0 trips day⁻¹ and were positively correlated with day length. Early winter foraging trips more closely matched day length than late winter foraging trips. The data suggest that individuals maximize foraging time during the early winter period, likely to recover body mass following the breeding season and molt. The more attenuated response of foraging trip durations to increasing day length in late winter may be related to differences in local resource availability or individual behaviors prior to the upcoming breeding season. Minimum food requirements also exhibited a seasonal cycle with a mid-winter minimum. On average, minimum food requirements were estimated at 0.70 ± 0.12 kg day⁻¹. Extrapolated to the regional population of gentoo penguins, winter food requirements by gentoo penguins were equivalent to roughly 33% of annual krill catches by commercial fisheries in the South Shetland Island region over the past decade. Current expansion of the gentoo population and the krill fishery in the southern Scotia Sea warrants continued monitoring of gentoo penguins during winter.     

Foraging behaviour and feeding locations of Rock Shags Phalacrocorax magellanicus from a colony in Patagonia, Argentina

During 1996 and 1997, foraging Rock Shags Phalacrocorax magellanicus were studied at Punta Loma, Argentina using radio-transmitters deployed on ten adult shags during the chick-rearing period. Rock Shags undertook 2.6 ± 0.6 sd trips per day. The mean duration of a feeding trip was 2.6 ± 0.7 hours. A bird spent 36% of daylight hours away from the colony on feeding trips, diving for 92% of the foraging trip, and made a mean of 106 dives per foraging trip. Foraging trip duration was strongly correlated with the total number of dives made in one foraging trip. Rock Shags fed mainly in water less than 10m deep with a gravelly sand bottom and within 5 km of shore. Mean foraging range was 3.8 ± 2.6 km and 2.6 ± 2.3 km for 1996 and 1997, respectively. These results suggest a high foraging effort (diving time per foraging trip) for Rock Shags, presumably associated with poor food conditions close to the colony. Comparison is made with other Phalacrocorax species.     

Patch choice and risk: relative competitive ability is context dependent

The relative abilities of individual cichlids, Tilapia zillii to obtain food under scramble competition was highly repeatable between trials using a single input source, regardless of whether the input was constant or variable. However, when given a choice between two patches differing only in their temporal variability in input about an identical mean, an individual's rank based on intake in one patch was uncorrelated with either its intake in the other patch or its intake in the single-patch trials. In the two-patch trials, certain individuals both spent more time in food patches and visited patches more often than others, and overall the fish spent more time in the constant rate patch than the variable patch, leading to more items being consumed from the constant rate patch. We discuss possible causes and consequences of this dependence of relative competitive ability on the context of the foraging situation. Copyright 1999 The Association for the Study of Animal Behaviour.     

Adjustments in the Time,Distance and Direction of Foraging in Dinoponera quadriceps Workers

We measured individual decisions regarding the adjustments of time, distance and direction of foraging in Dinoponera quadriceps. We observed two colonies in an area of secondary Atlantic Forest, FLONA-ICMBio, in Northeastern Brazil. The workers were individually marked. We recorded the displacement of workers, their returns to the nest with and without food, the time spent searching for food, maximum and total distance, inter-trip latency and direction of trips. The time spent searching for food, maximum distance and transport velocity did not vary with food size. The previous trip success reduced the latency between foraging trips and increased the percentage of success on the next trip. However, this previous success did not demonstrate a significant variation relative to the time spent searching on the next trip or direction of search. The workers maintained an individual directional fidelity during foraging. The adjustments of these foraging variables under individual control contributed to the efficiency at the colony level. D. quadriceps is compatible with the central place theory and risk sensitivity model of behavior.     

The individual counts: within sex differences in foraging strategies are as important as sex‐specific differences in masked boobies Sula dactylatra

Understanding how animals allocate their foraging time is a central question in behavioural ecology. Intrinsic factors, such as body mass and size differences between sexes or species, influence animals’ foraging behaviour, but studies investigating the effects of individual differences in body mass and size within the same sex are scarce. We investigated this in chick‐rearing masked boobies Sula dactylatra, a species with reversed sexual dimorphism, through the simultaneous deployment of GPS and depth‐acceleration loggers to obtain information on foraging movements and activity patterns. Heavier females performed shorter trips closer to the colony than lighter females. During these shorter trips, heavier females spent higher proportions of their flight time flapping and less time resting on the water than lighter females did during longer trips. In contrast, body mass did not affect trip duration of males, however heavier males spent less time flapping and more time resting on the water than lighter males. This may occur as a result of higher flight costs associated with body mass and allow conservation of energy during locomotion. Body size (i.e. wing length) had no effect on any of the foraging parameters. Dive depths and dive rates (dives h^?1) were not affected by body mass, but females dived significantly deeper than males, suggesting that other factors are important. Other studies demonstrated that females are the parent in charge of provisioning the chick, and maintain a flexible investment under regulation of their own body mass. Variation in trip length therefore seems to be triggered by body condition in females, but not in males. Consequently, shorter trips are presumably used to provision the chick, while longer trips are for self‐maintenance. Our findings underline the importance of accounting for the effects of body mass differences within the same sex, if sex‐specific foraging parameters in dimorphic species are being investigated.     

Time constraint on food choice in provisioning blue tits, Parus caeruleus: the relationship between feeding rate and prey size

Previous work on food-provisioning behaviour in blue tits suggested that the parents could gather larger prey items only by making longer foraging excursions, for example, by being more selective or by reaching more distant (and less exploited) feeding sites. Here, I show that within-nest, within-day variation in size of prey delivered by the parent could be explained by the time since its last visit. In unmanipulated conditions, size of larvae tended to increase with the time spent away from the nest. A significant positive relationship was more likely at high provisioning rates, suggesting that periods of intense feeding limited the size of prey delivered to the brood. To assess the effect of less intense feeding on prey size, I experimentally increased food availability to the tits. The parents could decide whether to eat the extra food or feed it to the nestlings. In both cases, food supplementation could result in longer time lags between natural feedings. Food-supplemented parents consumed the extra food and fed it to their nestlings, made longer foraging trips and delivered larger natural larvae than controls. In this group, size of larvae was more constant during the observation period and was independent of the time since the parent's last visit. This suggests that, below some value of visit rate, prey size is no longer limited by the duration of the foraging trip. The results support the view that tits continually vary visit rate and prey size. There is some evidence that these adjustments are made by changing food selectivity in response to changes in the state of the brood and of the parents.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Testing assumptions of central place foraging theory: a study of Adélie penguins Pygoscelis adeliae in the Ross Sea

We investigated central place foraging (CPF) in the context of optimal foraging theory in Adélie penguins Pygoscelis adeliae of the southern Ross Sea by using satellite tracking and time‐depth recorders to explore foraging at two spatio‐temporal scales: within the day‐to‐day (sub‐mesoscale: single foraging trip, 10s of km²) and the entire breeding season (mesoscale: trips by multiple individuals across the collective foraging area, 100s of km²). Specifically, we examine whether three basic assumptions of the Orians–Pearson CPF model, shown to occur in other CPF species, are met: 1) within a patch, the rate of prey acquisition declines with time spent in that patch; 2) food is distributed in discrete patches and is not available between those patches; and 3) CPF species have knowledge of the potential (or average, at least) feeding rate within their universe of patches, and use this knowledge to determine their foraging strategy when planning or engaging in a foraging trip. We found that prey consumption rates did not decline with time spent in patches, and penguins foraged to some degree most of the time when at sea. Food availability, as measured by foraging dive rate, appeared to be predictable within the same day at the same location, but predictability broke down after 2 d at distances > 10 km away. We conclude that the assumptions of the Orians–Pearson CPF model are not a good fit to the circumstances of Ross Sea penguins, which clearly are central place foragers.     

Age-related changes in foraging in the German cockroach (Dictyoptera: Blattellidae)

The dynamics of exploitation of standard experimental food sources by the German cockroach, Blattella germanicaL. (Blattellidae), were analyzed in an urban habitat in relation to developmental stage. The data presented here stress differences in foraging capacities between small (first-and second-instar) larvae and animals of other developmental stages. The first animals to arrive in a food patch presented a developmental-stage distribution significantly different from that of the general population. Adults and large larvae (fifth and sixth instars) were the first to find food sources and, in particular, before small larvae. Significant differences appeared between developmental stages concerning givingup time and the time animals left a patch. Small larvae were significantly underrepresented in a patch just before food exhaustion but they were significantly more numerous than expected just after depletion. Small larvae remained in the vicinity of a depleted food dish longer than animals of other developmental stages. Adults left patches as soon as these were depleted, long before small larvae did. Developmental stage influenced rate of departure. These observations indicate that cockroaches improve their foraging performance as they grow larger.     

Relationships between brood size and parental provisioning performance in chinstrap penguins during the chick guard phase

 Chinstrap penguins (Pygoscelis antarctica) normally lay two eggs, but brood size is often reduced by mortality during incubation or after hatching. We hypothesized that this variation in brood size would affect the parents’ foraging behavior and their chick provisioning performance. We studied patterns of adult foraging trip duration and frequency, food load delivery, and chick growth rates in relation to brood size during the guard phase in four breeding seasons (1991–1994) on Seal Island, Antarctica. Within a given year, parents with two chicks made more frequent foraging trips to sea and may have transported larger food loads to the nest; however, the duration of foraging trips was unrelated to brood size. Overall, parents with two chicks spent ∼15% more time at sea than parents with only one chick. Both the frequency and duration of foraging trips varied between years. Foraging trip duration may partly reflect the birds’ foraging radius, which probably varies with time in response to shifts in krill distribution. Chick growth rate varied betwen years, but was related to brood size only in 1992, when chicks from two-chick broods grew significantly more slowly than chicks from one-chick broods. Food loads transported to chicks, as well as chick growth rates, were highest in 1994, when concurrent hydroacoustic studies indicated that regional krill biomass was severely depressed. This apparent anomaly suggests that the spatial scale of the krill survey may have been too coarse to detect some high-density krill aggregations within the penguins’ foraging range. Received: 26 September 1995 / Accepted: 12 May 1996     

Overnight foraging trips by chick‐rearing Nazca Boobies Sula granti and the risk of attack by predatory fish

Most tropical booby species complete breeding foraging trips within daylight hours, thus avoiding nights at sea. Nazca Boobies Sula granti are unusual in this respect, frequently spending one or more nights away from the nest. We used GPS dataloggers, time‐depth recorders, and changes in body weight to characterize foraging trips and to evaluate potential influences on the decisions of 64 adult Nazca Boobies to spend a night at sea, or to return to their chicks on Isla Española, Galápagos, in daylight hours. The tagged birds foraged east of Isla Española, undertaking both single‐day (2–15 h, 67% of trips) and overnight trips (28 h–7.2 days, 33%), and executing 1–19 foraging plunge‐dives per single‐day trip. Birds might forage longer if they are in nutritional stress when they depart, but body weight at departure was not correlated with trip length. Birds might be expected to return from longer trips with more prey for young, but they returned from single‐day and overnight trips with similar body weights, consistent with previous indications that Nazca Boobies forage until accumulating a target value of prey weight. Birds with a lower dive frequency during the first 5 h of a trip were more likely to spend the night at sea, suggesting that they might choose to spend the night at sea if prey capture success was low. At night, birds almost never dived and spent most of their time resting on the water’s surface (11.8–12.1 h, > 99% of the time between civil sunset and civil dawn). Thus, the night is an unproductive time spent among subsurface predators under low illumination. The birds’ webbed feet provided evidence of this risk: 24% of birds were missing > 25% of their foot tissue, probably due to attacks by predatory fish, and the amount of foot tissue lost increased with age, consistent with a cumulative risk across the lifespan. In contrast, other tropical boobies (Blue‐footed Sula nebouxii and Brown Boobies Sula leucogaster), which do not spend the night on the water, showed no such damage. These results suggest that chick‐rearing Nazca Boobies accept nocturnal predation risk on occasions of low prey encounter during a foraging trip’s first day.     

VARIATION IN THE FORAGING LOCATION OF ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA) AND THE EFFECTS ON DIVING BEHAVIOR

This study investigated how female Antarctic fur seals adapt their foraging behavior, over time scales of days, to spatial unpredictability in the distribution of their food. Lactating Antarctic fur seals are central-place foragers that feed on highly patchy but spatially and temporally dynamic food. We measured the foraging distribution of 28 fur seals to test whether variation in foraging trip durations was reflected in variation in the location of foraging and the diving behavior of seals at sea. Based on the maximum distance travelled from the breeding beach, three categories of foraging trips were denned: those to the continental shelf area ( n = 12, median = 71 km), to oceanic water ( n = 11, median =164 km), and to farther offshore oceanic waters ( n = 5, median = 260 km). Trip duration and mean surface speed were positively correlated with the maximum distance travelled from the breeding beach. Seals on longer trips spent proportionally less of their time submerged, but there was no significant difference in the total number of dives or the total time spent foraging by seals in relation to trip duration. Evidence from this study and previous work investigating energy gain suggests that an animal on a longer foraging trip could potentially have a higher mean energy return per dive than a similar animal on a shorter foraging trip. Evidence presented suggests that the type of foraging trip (near or far) is not predetermined by the animal but may be a simple response to the stochastic distribution of the resources available.     

The Effect of Feeding History on Prey Capture Behaviour in the Orbweb Spider Argiope keyserlingi Karsch (Araneae: Araneidae)

The prey capture behaviour of the orb-web spider Argiope keyserlingi Karsch was examined experimentally by subjecting spiders to two different feeding regimes (food deprived and food satiated) and three types of prey: Drosophila, blowflies (Lucilia cuprina) and bees (Apis mellifera). The attack behaviour of the spiders was influenced by both their foraging history and the type of prey. Food deprived spiders attacked Drosophila and bees more frequently than food satiated spiders, and food satiated spiders travelled more slowly to any of the prey types than food deprived spiders. Furthermore, Drosophila were never wrapped in silk but only grasped with the chelicerae, whereas both blowflies and bees were always wrapped. This provides experimental confirmation that feeding history affects the decision of orb-web spiders to accept or reject any given prey.     

Does the foraging strategy of adult Short‐tailed Shearwaters cause obesity in their chicks?

The chick provisioning behaviour of Short-tailed Shearwaters Puffinus tenuirostris breeding at the northern edge of their distribution on Montague Island, New South Wales, was examined in February and March 1997. The duration of individual foraging trips of parents, weight changes of adults and chicks, and meal sizes delivered to chicks were determined. It was found that individual parents mixed a long foraging trip to Antarctic waters (14.4±2.0 days) with one to three short foraging trips (1.36±0.7 days, mode=1 day). Adults gained body mass on long trips and lost weight on short trips. The size of meals fed to the chicks was significantly greater after a long trip (161±21 g) than after a short trip (135±28 g), although short trips increased the overall chick feeding frequency. The variable number of short trips made by adult Short-tailed Shearwaters and the relationship between short trips and adult body condition were consistent with current life-history theory: adults do not sacrifice their own body condition to increase food delivered to their chicks.
Modelling revealed that this dual foraging strategy inevitable leads to chicks enduring long intervals between meals. These long intervals may have led to the evolution of an over-feeding strategy by parents and the nestling obesity reported in this shearwater. The durations of the long trips from Montague Island were significantly greater than those for Short-tailed Shearwaters breeding at the centre of their distribution in Tasmania, although there was no significant difference in the length of short trips. A commitment to feed regularly in Antarctic waters may explain why the breeding distribution of this species does not extend much further north.     

Patterns of energy acquisition by a central place forager: benefits of alternating short and long foraging trips

In some seabirds, foraging trips have been defined as eitherlong or short, with the length of time spent traveling to theforaging area apparently a critical feature in determining foragingtrip length. Using logger technology, together with complimentarydata from published studies, we investigated traveling and foragingtimes in 18 free-living Adélie Penguins Pygoscelis adeliae,which were foraging for chicks. Most deep, foraging dives weredistributed around the center of the foraging trip. This centraltendency was particularly apparent if the cumulative amountof undulations in the depth profile (indicative of prey capture)was considered during deep dives; values started to increasebefore 20.9% and ceased after 67.2% of the dives had occurred.This concentration of the feeding activity in the middle ofthe foraging trip indicates that birds traveled to and froma prey patch whose location varied little over the birds' trips.These data form the basis for a simple model that uses travelingand foraging times together with projected rates of prey ingestionand chick and adult gastric emptying to determine that thereare occasions when, to optimize rates of prey ingestion whileat sea for both adults and chicks, birds should conduct foragingtrips of bimodal lengths.     

Effects of Foraging Experiences on Residence Time of the Predatory Mite <Emphasis Type="BoldItalic">Neoseiulus womersleyi</Emphasis> in a Prey Patch

We investigated the effects of foraging experiences on the residence time of Neoseiulus womersleyi in a currently inhabited prey (Tetranychus urticae) patch. Satiated predators that had experienced starvation stayed longer in a current patch than those that had not experienced starvation. Satiated predators that had experienced a prey-rich patch showed approximately the same residence time in the current patch irrespective of the number of prey therein. By contrast, satiated predators that had experienced a prey-poor patch stayed longer in a current patch of high prey density than in one of low prey density. N. womersleyi appears to determine residence time in the current patch based on foraging experiences together with the quantity of prey in the current patch.     

Development of foraging behavior in juvenile northern elephant seals

Rapid development of foraging ability is critical for phocids. In northern elephant seals Mirounga angustirostris , juvenile survivorship is low compared with adults and foraging difficulties are potentially associated with increased mortality. At Año Nuevo, California, foraging behavior of nine juvenile females during their third foraging migration and five juvenile females on their fourth foraging migration were documented using a variety of commercially available and custom time depth recorders. Foraging success, diving ability, time at depth, bouts of behavior and body composition changes were compared between trips to sea. There were no significant differences in foraging success measured as mass gain between the third and fourth trips to sea. There were differences in how energy was deposited between lean and adipose tissue compartments. Diving ability developed between trips to sea, reflected in significant increases in depth, dive duration and bottom time. Development also occurred within trips to sea. Depth, dive duration and bottom time increased with time at sea. Aerobic capacity appears to increase between the third and fourth trip, with a significantly increased percentage of total time submerged and a significantly lower diving rate. All juveniles on the fourth trip and four out of nine juveniles on the third trip followed marked diel patterns, foraging deep during the day and shallow at night. Like adults, juveniles appeared to stay primarily aerobic with surface intervals independent of dive durations. These results confirm that female juvenile northern elephant seals undergo important developmental changes in foraging behavior between the third and fourth trip, but these changes do not significantly impact foraging success.