Analyses of Two Parasitoids with Convergent Foraging Strategies

We compared the foraging strategies of two key braconid endoparasitoids of the tobacco budworm (Heliothis virescens Fab.), Cardiochiles nigriceps Vier. and Microplitis croceipes Cresson, that differ in host and habitat range but otherwise share comparable, overlapping niches. The most important host-location cues by far for both species were materials associated with damaged plants. Both species demonstrated a significant preference for volatiles released from plants damaged by H. virescens larvae over those released from undamaged tobacco and cotton plants. In choice experiments with damaged tobacco versus cotton, M. croceipes showed a significant preference for cotton plants. In contrast, C. nigriceps preferred damaged tobacco plants. Plant compounds provoked a strong response even when released from systemically induced plants (from which damaged leaves, host, and host by-products were removed). C. nigriceps appears to have a much keener ability to locate hosts over long distances than M. croceipes. This observation may be related to the highly specialized nature of this parasitoid. The possible adaptive significance of the foraging behaviors of these two parasitoids is discussed.     

Seasonal patterns of diet and ranging in two species of tamarin monkeys: Stability versus variability

From June through December, data were collected on the diet and ranging patterns of moustached (Saguinus mystax) and saddle-back (Saguinus fuscicollis) tamarin monkeys in the Amazon Basin of northeastern Peru. During this 7-month period, insects and nonleguminous fruits accounted for 83% of tamarin feeding and foraging time. Despite marked seasonal variation in rainfall and forest productivity, patterns of habitat utilization, day range, dietary diversity, resource exploitation, and activity budget remained relatively stable throughout the year. Moustached and saddle-back tamarins appear to solve problems of food acquisition and exploit patchily distributed feeding sites using a relatively limited set of foraging patterns. In general, these primates concentrate their daily feeding efforts on several trees from a small number of target plant species. These feeding sites are uncommon, produce only a small amount of ripe fruit each day, and are characterized by a high degree of intraspecific fruiting and flowering synchrony. Trees of the same species are frequently visited in succession, and individual feeding sites are revisited several times over the course of 1–2 weeks. This type of foraging pattern occurred during both dry and wet seasons and when exploiting fruit, nectar, legume, and exudate resources. Seasonal variation in the percentage of feeding and foraging time devoted to insectivory was also limited. In this investigation, there was no consistent evidence that temporal changes in overall forest fruit production had a major impact on the feeding, foraging, or ranging behavior of either tamarin species.     

Effects of Hunger on Foraging Behavior of Age-0 Smallmouth Bass in a Heterogeneous Environment

We examined how hunger affected habitat use by juvenile smallmouth bass, Micropterus dolomieu, as they moved among a variety of habitat patches. Hungry and satiated fish were placed in an artificial stream that contained three types of habitat patches: pools with uniform depth and low water velocities, mazes with uniform depth and moderate water velocities, and riffle-pool complexes with varying depth and the highest water velocities. Food was only available in the riffle-pool complexes. Hungry fish spent more time in the riffle-pool complex than satiated fish did. However, hungry and satiated fish did not differ in the time it took to exit the pool they were initially placed in, the number of patches entered, or the number of times they moved among patches. Both hungry and satiated fish frequently entered other patches after foraging successfully in the riffle-pool complexes. There was wide variation in foraging behavior among individuals in both treatment groups, and we consistently observed individuals that did not alter their foraging behavior in response to the difference in food availability among patches.     

Costs and Benefits of Food Foraging for a Braconid Parasitoid

The costs and benefits of food foraging for the insect parasitoid, Phanerotoma franklini Gahan (Hymenoptera: Braconidae), were examined. Feeding benefited P. franklini by increasing longevity, but did not influence egg maturation. Costs of feeding are represented by the time required to locate and consume food. Field observations of starved wasps revealed that wasps spent approximately 25% of their time grazing on substances on the surface of cranberry foliage. However, our initial laboratory tests to determine the value of these substances were inconclusive. We also examined how grazing behavior integrated with host foraging by comparing the attack rate of fed and unfed wasps held under different host densities. While feeding status did not affect the mean attack rate of wasps, starvation enhanced the probability that a wasp will engage in movement over the host plant foliage. We conclude that feeding benefits P. franklini by increasing longevity, that travel time to food sources is likely to be low, and that the observed food-foraging behavior does not appear to influence host-foraging efficiency.     

Foraging strategies in solitary parasitoids: The trade-off between female and offspring mortality risks

Summary It is often assumed that oviposition rate is the currency that parasitoids should maximize in order to maximize reproductive success. Female parasitoids foraging in a patchy environment face a variety of mortality risks that influence the survival of both themselves and their offspring. Maximizing oviposition rate ignores these risks. A model is developed to analyse the influence of female and offspring mortality risks on optimal patch residence time in time-limited solitary parasitoids. The optimal compromize between minimizing a female's own mortality risks and the mortality risks of her offspring in characterized. The optimal patch residence time is shown to be dependent on the relative magnitude of these mortality risks, as well as the rate with which reproductive success accumulates while on a patch. If travel time between patches is not fixed but a random variable, the optimal patch residence time decreases. However, variability in travel time increases expectations of total reproductive success. The model is illustrated with a case study in two aphid parasitoids.     

Modeling monkeys: A comparison of computer-generated and naturally occurring foraging patterns in two species of neotropical primates

We present a series of computer-generated foraging models (random movement, olfactory navigation, and spatial memory) designed to examine the manner in which sensory cues and cognitive skills might be used by rainforest monkeys to locate patchily distributed feeding sites. These simulations are compared with data collected in the Amazon Basin of northeastern Peru on the foraging patterns of two species of neotropical primates, the moustached tamarin monkey (Saguinus mystax) and the saddle-back tamarin monkey (Saguinus fuscicollis). The results indicate that, although tamarins may rely on olfactory cues to locate nearby feeding sites, their foraging patterns are better explained by an ability to maintain a detailed spatial map of the location and distribution of hundreds of feeding trees in their home range. There is evidence that such informationis retained for a period of at least several weeks and is used to minimize the distance traveled between widely scattered feeding sites. The use of computer simulations provides a powerful research tool for generating predictive models regarding the role of memory and sensory cues in animal foraging patterns.     

Tamarin polyspecific associations: Forest utilization and stability of mixed-species groups

Niche separation is likely to play a key role in the formation of mixed-species groups. Saddle-backed tamarins (Saguinus fuscicollis) were studied at three sites with different primate communities in northern Bolivia: (1) with red-bellied tamarins,S. labiatus; (2) with emperor tamarins,S. imperator; and (3) without a congeneric species. The degree of association is higher betweenS. labiatus andS. fuscicollis than betweenS. imperator andS. fuscicollis and is related to differences in forest utilization between associating pairs. Niche separation is found to be greater betweenS. labiatus andS. fuscicollis than betweenS. fuscicollis andS. imperator. The mean height and habitat utilization ofS. fuscicollis does not differ greatly across the three sites, nor does the height of tamarins in and out of association. It is concluded that combined with differences in body size and dietary overlap, vertical segregation plays an important role in tamarin polyspecific associations (increasing the potential of both foraging and anti-predatory benefits) and that this is not a consequence of vertical displacement ofS. fuscicollis by its dominant congeners.     

Chuck Vocalizations of Wild Female Squirrel Monkeys (Saimiri sciureus) Contain Information on Caller Identity and Foraging Activity

Analysis of the acoustic signal of the chuck vocalizations of adult female squirrel monkeys (Saimiri sciureus) in Parque Nacional de Manu, Peru, revealed consistent differences within and between individuals. We quantified four peak frequency parameters: (a) the peak frequency of single chucks, (b) the first and (c) the second peak frequencies of double chucks, and (d) the peak difference: the difference between the first and the second double chuck peaks. One-way ANOVAs and a posteriori comparisons of these variables revealed that each distinguished more than 70% of all possible pairs of females. When all double chuck measures were included in a discriminant analysis, 57% of double chucks were correctly assigned to the caller. Another category of information potentially encoded in the acoustic structure of chuck vocalizations is foraging activity. When the chucks of squirrel monkeys during foraging and nonforaging activities were compared, the single chuck peak frequency, and the first peak frequency and the peak difference of double chucks, were significantly reduced during foraging contexts. Previously Boinski and Mitchell (1992) concluded that chucks facilitate group cohesion among widely dispersed troop members by providing information of the location of callers; the rate of chucks produced by an adult female increases as she becomes more spatially and visually separated from other adult females. The additional information potentially conveyed by chucks on caller identity and foraging activity documented in these new analyses further emphasizes the role chucks serve to enhance group coordination and cohesion.     

Foraging behavior ofPachydiplax longipennis (Odonata: Libellulidae)

Food intake, prey availability, and prey capture behavior at feeding areas were quantified in the dragonflyPachydiplax longipennis by observing focal individuals on artificial perches, where they exhibited marked short-term site fidelity. Prey capture success was high and relatively constant, but the frequency and duration of feeding flights depended on the time of day and season (at least in part because of associated variation in temperature and other physical factors), study site, sex, density of other dragonflies, and prey density. Individuals rapidly responded temporally and spatially to changes in prey availability, particularly to localized prey concentrations.     

Foraging Efficacy of a Larval Parasitoid in a Cotton Patch: Influence of Chemical Cues and Learning

Plant-herbivore chemical signals and behavioral plasticity may enhance parasitoid host-foraging efficacy in the field; however, no studies have quantified the potential benefits from these factors under field-type conditions. The effect of plant-herbivore signals and learning on the foraging efficacy of Microplitis croceipes was quantified by directly observing and recording total and sequential duration of various foraging behaviors relative to 5 randomly placed herbivore-damaged and host-infested cotton plants and 20 undamaged and non-host-infested plants. Microplitis croceipes spent significantly more time searching (flying and antennation) on host infested versus uninfested plants. Antennation time was significantly and negatively correlated with successive host stings. Contrary to expectations of increased duration, flight time remained constant throughout the foraging bout, which may indicate that there was some learning associated with flight. These results suggest that plant-herbivore chemical signals and learning enhances the foraging efficacy of M. croceipes.     

Settling Decisions and Heterospecific Social Information Use in Shrikes

Animals often settle near competitors, a behavior known as social attraction, which belies standard habitat selection theory. Two hypotheses account for these observations: individuals obtain Allee benefits mediated by the physical presence of a competitor, or they use successfully settled individual as a source of information indicating the location of high quality habitat. We evaluated these hypotheses experimentally in two species of shrikes. These passerine birds with a raptor-like mode of life impale prey to create larders that serve as an indicator of male/habitat quality. Thus, two forms of indirect information are available in our system: a successfully settled shrike and its larder. Typically these two cues are associated with each other, however, our experimental treatment created an unnatural situation by disassociating them. We manipulated the presence of larders of great grey shrikes and examined the settling decisions of red-backed shrikes within and outside the great grey shrike territories. Male red-backed shrikes did not settle sooner on plots with great grey shrikes compared to plots that only contained artificial larders indicating that red-backed shrikes do not use the physical presence of a great grey shrike when making settling decisions which is inconsistent with the Allee effect hypothesis. In contrast, for all plots without great grey shrikes, red-backed shrikes settled, paired and laid clutches sooner on plots with larders compared to plots without larders. We conclude that red-backed shrikes use larders of great grey shrikes as a cue to rapidly assess habitat quality.     

Foraging behaviour of planktivorous fish in artificial vegetation: the effects on swimming and feeding

In the littoral zones of lakes, aquatic macrophytes produce considerable structural variation that can provide protection to prey communities by hindering predator foraging activity. The swimming and feeding behaviour of a planktivore, Pseudorasbora parva(Cyprinidae) on its prey (Daphnia pulex) was studied in a series of laboratory experiments with varying densities (0, 350, 700, 1400, 2100 and 2800 stems m^–2) of simulated submerged vegetation. Prey availability was varied from 0.5, 1.0, 2.0, 5.0, 10.0 and 25.0 prey l^–1. As the stem density increased, the predator's swimming speed and the number of prey captured decreased relative to feeding in open water. A good relation existed between the number of successful prey captures and swimming speed with the average stem distance to fish body length ratio (D). An abrupt reduction in feeding and swimming was recorded when D was reduced to values less than one.     

Foraging and building in subterranean termites: task switchers or reserve labourers?

Task-switching between foraging and building in workers of Nasutitermes exitiosus (Termitidae), a subterranean termite, was followed using mark-recapture. Foragers were collected from wood-filled drums and marked with Nile blue (ca. 12,500 per colony), whereas builders were collected from damaged sections of mounds and marked with Neutral red (ca. 18,500 per colony). Two protocols were followed: the first marked foragers and then damaged the mound; the second reversed this order. In the first protocol few foragers switched to building, i.e. blue-marked workers in mound samples, measured either as numbers (81 ± 28) or proportion of originally marked (0.66 ± 0.20 %); whereas more than double blue-marked foragers remained foraging (222 ± 64 workers or 1.76 ± 0.44 %). In the second protocol few builders had switched to foraging, i.e. red-marked workers found in the first drum sample (82 ± 33 workers or 0.29 ± 0.09 %). Up to eight drums were sampled for foragers over about 80 days; the numbers of blue-marked workers decreased slowly (ca. 1.6 workers per day) whereas those of red-marked workers increased slowly (ca. 2.4 workers per day). These results suggest that relatively few termite workers switch directly between foraging and building. The possibility of builders being recruited from other tasks is discussed. Received 24 March 2005; revised 27 June 2005; accepted 2 July 2005.     

Foraging behavior of the communal spider,Philoponella republicana (Araneae: Uloboridae)

The communal orb-weaving spider, Philoponella republicana,was observed in the subtropical moist forest of Southeast Peru. These spiders live in colonies of conspecifics whose individual orbs are connected by silk. The wrapping of a prey prior to feeding is a large component of the prey capture process because P. republicanahas no venom with which to kill an insect. Wrapping time was the only aspect of prey capture that was strongly correlated with the size of the insect captured. Occasionally we observed several individuals working together to wrap a prey item. These joint efforts were more frequent on prey larger than the capturing spider. Although group captures accounted for only 5.5% of captures, they represented 14.7% of the biomass obtained. A comparison of the relationship between wrapping time and prey size for solitary and group efforts suggested that, by working together, the spiders reduced their total handling time. In most cases only one spider fed on the captured prey.     

Social Learning of a Novel Foraging Task by Big Brown Bats (Eptesicus fuscus)

Acquiring information via observation of others can be an efficient way to respond to changing situations or learn skills, particularly for inexperienced individuals. Many bat species are gregarious, yet few studies have investigated their capacity for learning from conspecifics. We tested whether big brown bats (Eptesicus fuscus) can learn a novel foraging task by interacting with knowledgeable conspecifics. In experimental trials 11 na?ve bats (7 juveniles, 4 adults) interacted freely with trained bats that were capturing tethered mealworms, while in control trials 11 na?ve bats (7 juveniles, 4 adults) flew with untrained bats. Na?ve bats were then assessed for their ability to capture tethered mealworms. While no bat in the control group learned the task, a significant number of experimental bats, including juveniles with little or no experience foraging, showed evidence of learning. Eighty-two per cent of experimental bats and 27% of control bats directed feeding buzzes (echolocation calls associated with prey capture) at the mealworm. Furthermore, seven experimental bats (64%) showed evidence of learning by attacking and/or capturing the mealworm, while no bat in the control group attacked or captured the prey. Analyses of high-speed stereo video recordings revealed increased interaction with demonstrators among bats attacking or capturing the mealworm. At the time they displayed evidence of learning, bats flew closer together during feeding buzzes than during other portions of trials. Our results demonstrate that social interaction with experienced bats, and listening to feeding buzzes in particular, may play an integral role in development of foraging skills in bats.     

Home Range Size and Habitat Use in the Black Lion Tamarin (Leontopithecus chrysopygus)

The black lion tamarin (Leontopithecus chrysopygus) is a threatened species due to past destruction of its natural habitat, thus knowledge of its home range size and use is essential in planning for its conservation. I studied one group for 4 months in the Morro do Diabo State Park, in São Paulo State, Brazil. I estimated the home range of the group to be 64 and 127 ha via the quadrat and convex polygon methods, respectively, while a composite method yielded an estimate of 106 ha. They ate insects most frequently (38% of scans), a resource which was distributed throughout their home range. Fruit was the second major resource, but when it was not available, they ate more gum. The exploitation of fruits was associated with dryland forest, while gum-feeding occurred mainly in swamp forest. The study group used a transition zone between dryland and swamp forest most frequently, and all of their sleeping trees were located there. Although the vegetation reached 15–20 m, the group spent most of the time (55%) in the upper understory, between 4 and 8 m high. The study group revealed more specific habitat needs than thought previously, suggesting that the current population may be smaller than estimated.     

Foraging Area Marking in Two Related Tetramorium Ant Species (Hymenoptera: Formicidae)

The related ants Tetramorium caespitum and T. impurum mark their foraging area in a species-specific, home range and short-lasting manner. Indeed, ants reaching a new area have a slow linear speed which increases during the marking. Conspecific ants are arrested and attracted by marked areas, while heterospecific ants are reluctant to visit them. However, when the latter do visit marked areas, they move more quickly and less sinuously than conspecific ants and do not stay on the areas. The marking is performed in about 3 min by T. caespitum and in 3 to 6 min by T. impurum. If not reinforced, the marking vanishes in the same time intervals. Neither poison gland nor last sternite extracts reproduce the activity of naturally marked areas, whereas a Dufour gland extract does exactly that. Foraging ants touch the ground with the tip of their gaster. Consequently, we can postulate that the workers mark their foraging area with the contents of this gland, which is associated with the sting apparatus, and that they deposit with the extremity of the gaster. Alien conspecific ants are seldom aggressive to one another, even on marked areas. When encountering each other on unmarked areas, heterospecific ants present some aggressive reactions. On marked areas, their aggressiveness is enhanced and intruder ants are restless, while resident ones walk freely. On ground marked by T. impurum, ants of this species are more aggressive than antagonistic T. caespitum workers. The marking of foraging areas thus induces defense against heterospecifics but not against conspecific ants.     

Foraging behavior of redheads (Aythya americana) wintering in Texas and Louisiana

Redheads, Aythya americana, concentrate in large numbers annually in traditional wintering areas along the western and northern rim of the Gulf of Mexico. Two of these areas are the Laguna Madre of Texas and Chandeleur Sound of Louisiana. We collected data on 54,340 activities from 103 redhead flocks in Texas and 51,650 activities from 57 redhead flocks in Louisiana. Males and females fed similarly, differing neither in levels of feeding (percent of all birds in flock that were feeding) (p>0.90) nor in percentages of birds feeding by diving, tipping, dipping, or gleaning from the surface (p>0.10). The foraging level of redheads in the upper Laguna Madre region was relatively constant throughout two winters. Foraging of redheads in early winter in Louisiana was significantly greater than redhead foraging in the upper Laguna Madre, but by late winter, foraging by redheads in Louisiana had declined to the same level as that shown by redheads foraging in the upper Laguna Madre. The overall foraging level of redheads from Chandeleur Sound was greater (41%) than that of redheads in the upper Laguna Madre (26%), yet it was quite similar to the 46% foraging level reported for redheads from the lower Laguna Madre. Redheads in the upper Laguna Madre region of Texas fed more by diving than did those in the Chandeleur Sound and the lower Laguna Madre. Diving increased in frequency in late winter. Greater reliance by redheads on diving in January and February indicates that the birds altered their foraging to feed in deeper water, suggesting that the large concentrations of redheads staging at this time for spring migration may have displaced some birds to alternative foraging sites. Our results imply that the most likely period for food resources to become limiting for wintering redheads is when they are staging in late winter.     

Foraging strategy in a subterranean rodent,Spalax ehrenbergi: a test case for optimal foraging theory

Summary The foraging behavior of the subterranean mole rat Spalax ehrenbergi (Rodentia, Spalacidae) was tested according to the framework of optimal foraging theory. We compared the frequencies of food species hoarded in storage chambers of mole rats with the frequencies of these species occurring in the vicinity of the mole rats' nest mounds during the winter and spring seasons. In addition, we examined the food composition of several summer nest mounds. Laboratory observations were conducted in order to test the foraging behavior of mole rats under simulated subterranean conditions. The mole rat is a generalist and collects a variety of food species. Out of 33 plant species that were hoarded by mole rats in the 21 studied nest mounds, 61% (n=20) were geophytes, 21% (n=7) perennial herbs, 15% (n=5) annual herbs and 3% (n=1) dwarf shrubs. The frequency of each collected species in the 16 winter and spring nest mounds is in general accordance with its frequency in the mole rat's territory. This implies that the mole rat randomly samples the food reserve of its territory without special preference or directed search for a particular species. The collection or avoidance of any food item is not dependent on the presence or absence of any other food item. We suggest that the foragin generalism of the mole rat is a product of the constraints of a subterranean niche — the necessity to hoard food as much as possible in a limited time period and the high energetic investment of tunneling to the food items.