Time to revisit? A predator's previous successes and failures in prey capture determine its return time to patches.

In a heterogeneous environment containing multiple patches that may deplete and renew, a forager should be able to detect the quality of food resources within and among patches and choose to exploit them to best maximize returns. From the predator's perspective, the behavioral responses of the prey in a patch will be perceived as depletion when they retreat to refuge and renewal when they reemerge. A predator encountering responsive prey should manage predation risk, and thus behavioral resource depression, by optimally timing its return time to the patch based on prey behavior. We evaluated the foraging decisions of a predator that encountered patches differing in size of the refuge and prey density. We used little egrets and goldfish as predators and prey in an environment that contained three patches (pools). We manipulated prey density and refuge size and availability (using covers) and observed predator foraging behavior. When the egret had previously caught a fish it did not discriminate between the pools, and the return time was similar for all cover types. The fish densities also did not affect the egret decisions to return to pools. However, when it failed to catch fish, it returned sooner to the pool containing the small cover than the larger one. Additionally, after failing to catch fish in patches containing the highest prey density, the egrets subsequently preferred to return to such patches sooner. We show experimentally that previous failures influence the foraging decisions of a predator choosing how quickly to return to a previously visited patch.

Optimal foraging of little egrets and their prey in a foraging game in a patchy environment.

We explored the behavioral game between a predator, the little egret (Egretta garzetta), and a prey, the common goldfish (Carassius auratus), in a laboratory theater containing three fish pools. We tested the hypotheses that the egrets maximize their total capture success by responding to the fish's antipredatory behavior and that the behaviors of both players respond adaptively to the density distribution of fish among the pools. One experiment presented egrets with 15 fish per pool. The second experiment used a heterogeneous environment: pools 1, 2, and 3 had 10, 15, and 20 fish, respectively. Within each pool, fish could move between a safe, covered microhabitat and a risky, open microhabitat. Only the risky habitat had food, so fish were trading off food and safety by allocating the time spent in the two habitats. Egrets spent more total time in pools with more fish and returned to them sooner. Egrets maximized the number of fish they captured by following the matching rule of the ideal free distribution. The fish used the risky but productive habitat 65% of the time during experiments without egrets, but only 9% during experiments with 15 fish and egrets present somewhere in the theater. In addition, with egrets present, fish fine-tuned their behavior by reducing their use of the risky habitat as the egrets increased the frequency of their visits.

Age, intensity of infestation by flea parasites and body mass loss in a rodent host.

Parasitism by the flea Synosternus cleopatrae does not affect the body mass of its principal rodent host, Gerbillus andersoni under natural infestation levels. We hypothesized that the lack of negative effects of flea parasitism on rodent body mass could be related either to the low level of natural infestation or to the differential susceptibility of rodent age cohorts to flea parasitism. We tested these hypotheses by measuring body mass change under flea parasitism in (a) adult rodents infested with fleas above the natural infestation level (the first hypothesis) and (b) juvenile rodents infested with fleas at natural infestation levels (the second hypothesis). Adult individuals parasitized by a number of fleas higher than in nature lost body mass at higher rates than non-parasitized control individuals. Parasitism significantly affected daily body mass change of juvenile gerbils. Juvenile rodents parasitized by fleas at the natural level of infestation lost body mass faster and gained body mass slower than control animals. We suggest that some regulating mechanisms may limit natural flea densities at a point at which the negative effect on hosts is below the accuracy of our measurements. However, natural flea densities are sufficiently high to harm the more susceptible, juvenile cohort.

Age-biased parasitism and density-dependent distribution of fleas (Siphonaptera) on a desert rodent.

Parasites often confront conflicting demands when evaluating and distributing themselves among host individuals, in order to attain maximum reproductive success. We tested two alternative hypotheses about host preference by fleas in relation to the age of their rodent host. The first hypothesis suggests that fleas select adult over juvenile rodents because the latter represent a better nutritional resource (the "well-fed host" hypothesis), whereas the second hypothesis suggests that fleas prefer the weaker and less resistant juveniles because they are easier to colonise and exploit ("poorly fed host" hypothesis). We sampled fleas (Synosternus cleopatrae) on the gerbil (Gerbillus andersoni) in 23 different plots in the Negev desert and found an unequal distribution of fleas between adult and juvenile hosts. Furthermore, flea distribution changed as a function of flea density-from juvenile-biased flea parasitism (the "poorly fed host" hypothesis) at low densities to adult-biased flea parasitism (the "well-fed host" hypothesis) at high densities. Other factors that influenced flea preference were soil temperature and the presence of ticks. These results suggest that host selection is not an explicit alternative choice between adults and juveniles ("well-fed host" versus "poorly fed host" hypotheses), but rather a continuum where the distribution between adults and juveniles depends on host, parasite, and environmentally related factors.

The ecology of cutaneous leishmaniasis in Nizzana, Israel: infection patterns in the reservoir host, and epidemiological implications.

We conducted an extensive interdisciplinary study in an emerging focus of cutaneous leishmaniasis in the Western Negev Desert of Israel between July 1998 and February 2000. The aims of the this study were to determine (1) the reservoir hosts, (2) the distribution of the pathogen within the host range, (3) the associations of host, vector, and pathogen within defined habitats, (4) the demographic distribution of the pathogen within the host populations, and (5) to apply the newly acquired epizootiological data to explain morbidity patterns in humans. Fourteen square (60 m width) sampling plots were delimited in three types of habitats each with a different kind of substrate: loess, sand, and sand-loess ecotone. Rodents and sand flies were trapped and several environmental variables were measured. Leishmania infections in rodents were detected microscopically in stained smears of ear tissue and by a Leishmania-specific polymerase chain reaction. Results indicate that, contrary to previous reports, Psammomys obesus and not Meriones crassus is the main reservoir host in the region. Additional rodents (12 Gerbillus dasyurus and two M. crassus) were also found positive for Leishmania DNA. Prevalence of Leishmania infections amongst P. obesus was highest in loess habitats (65%), intermediate in the sandy-loess ecotone (20%), and 0% in the sandy habitats. Psammomys obesus individuals in the loess habitat of the Nizzana ruins were larger, on average (probably older), than those in the sandy habitat of the Mt. Keren junction. Sand fly density was positively correlated to soil moisture being higher in the relatively humid plots of Nizzana ruins and much lower in the drier sandy soil of Mt. Keren. Elucidation of fundamental ecological factors affecting this disease has helped explain an apparent discrepancy between the distribution of the disease in the zoonotic system and among humans.

Safety in numbers: sophisticated vigilance by Allenby's gerbil.

Since 1963, nonlinear predation theory has predicted that, at low population densities, victim species may well be mutualistic rather than competitive. Theory identifies this mutualism as a principal source of dynamic instability in the interaction. Using gerbils and trained barn owls, we conducted the first (to our knowledge) field tests of the theory's prediction of mutualism. The behavior of the gerbils confirms its existence.

The effect of barn owls (Tyto alba) on the activity and microhabitat selection of Gerbillus allenbyi and G. pyramidum.

Predation plays an important role in ecological communities by affecting prey behavior such as foraging and by physical removal of individual prey. In regard to foraging, animals such as desert rodents often balance conflicting demands for food and safety. This has been studied in the field by indirectly manipulating predatory risk through the alteration of cues associated with increased risk such as cover or illumination. It has also been studied by directly manipulating the presence of predators in aviaries. Here, we report on experiments in which we directly manipulated actual predatory risk to desert rodents in the field. We conducted a series of experiments in the field using a trained barn owl (Tyto alba) to investigate how two species of coexisting gerbils (Gerbillus allenbyi and G. pyramidum) respond to various cues of predatory risk in their natural environment. The gerbils responded to risk of predation, in the form of owl flights and owl hunger calls, by reducing their activity in the risky plot relative to the control plot. The strongest response was to owl flights and the weakest to recorded hunger calls of owls. Furthermore, when risk of predation was relatively high, as in the case with barn owl flights, both gerbil species mostly limited their activity to the safer bush microhabitat. The response of the gerbils to risk of predation disappeared very quickly following removal of the treatment, and the gerbils returned to normal levels of activity within the same night. The gerbils did not respond to experimental cues (alarm clock), the presence of the investigators, the presence of a quiet owl, and recorded "white noise". Using trained barn owls, we were able to effectively manipulate actual risk of predation to gerbils in natural habitats and to quantify how gerbils alter their behavior in order to balance conflicting demands of food and safety. The method allows assessment of aspects of behavior, population interactions, and community characteristics involving predation in natural habitats.

Tilman's predicted productivity-diversity relationship shown by desert rodents.

Tilman has developed a model to predict the number of plant species that can coexist competitively on a limited resource base. Species diversity first increases over low resource supplies, then declines as the environment becomes richer. Although Tilman 's model was developed to describe interspecific interactions between plant species, it may also apply to animal species. Tilman questions whether animals specialize on particular proportions of nutrients. However, we believe animals probably specialize on relatively subtle microhabitat differences, especially in a multispecies competitive regime. Thus, microhabitats may act like nutrients. We hypothesize that animal species, too, show a peaked curve of diversity over productivity. The present data provide a confirmation of the hypothesis using rodent species. We have investigated the number of rodent species along a geographical gradient of increasing rainfall. The gradient extends from extremely poor desert habitats to those with annual rainfall over 300 mm. Because of the aridity , precipitation reflects productivity. The diversity pattern in desert rodents agrees with that predicted by Tilman for plants. It even possesses similar asymmetry, rising steeply then falling slowly. The pattern is duplicated in rocky and sandy habitats, each of which has a distinct and almost nonoverlapping assemblage of species. As mean precipitation is closely correlated with the variability of precipitation, the diversity pattern might also be caused by a decline in the frequency of disturbances, models for which have been proposed by several investigators.

Habitat selection in two species of short-horned grasshoppers : The role of thermal and hydric stresses.

The roles of thermal and hydric stress in habitat selection by two species of short-horned grasshoppers was investigated through field and laboratory studies and computer simulation experiments. Psoloessa delicatula was found to possess an elaborate repertoire of thermoregulatory postures and shade seeking behavior. This grasshopper was found in habitats in the shortgrass prairie in which bare patches of ground were common. It was found to regulate its body temperature to a relatively constant level over the course of a sunny day. Eritettix simplex was found in patches of dense vegetation, and its body temperature was usually similar to ambient temperature. During controlled laboratory experiments, E. simplex nymphs lost water rapidly and thus were presumed to be restricted to more mesic environments where desiccation stress is less. A thermal energy budget model was constructed and showed that the habitat selected by P. delicatula would allow behavioral thermoregulation whereas the habitat selected by E. simplex should preclude regulation of body temperature by behavioral means.