Sounds of danger and post-traumatic stress responses in wild rodents: ecological validity of a translational model of post-traumatic stress disorder.

In the wild, animals face a highly variable world full of predators. Most predator attacks are unsuccessful, and the prey survives. According to the conventional perspective, the fear responses elicited by predators are acute and transient in nature. However, the long-term, non-lethal effects of predator exposure on prey behavioral stress sequelae, such as anxiety and post-traumatic symptoms, remain poorly understood. Most experiments on animal models of anxiety-related behavior or post-traumatic stress disorder have been carried out using commercial strains of rats and mice. A fundamental question is whether laboratory rodents appropriately express the behavioral responses of wild species in their natural environment; in other words, whether behavioral responses to stress observed in the laboratory can be generalized to natural behavior. To further elucidate the relative contributions of the natural selection pressures influences, this study investigated the bio-behavioral and morphological effects of auditory predator cues (owl territorial calls) in males and females of three wild rodent species in a laboratory set-up: Acomys cahirinus; Gerbillus henleyi; and Gerbillus gerbillus. Our results indicate that owl territorial calls elicited not only "fight or flight" behavioral responses but caused PTSD-like behavioral responses in wild rodents that have never encountered owls in nature and could cause, in some individuals, enduring physiological and morphological responses that parallel those seen in laboratory rodents or traumatized people. In all rodent species, the PTSD phenotype was characterized by a blunting of fecal cortisol metabolite response early after exposure and by a lower hypothalamic orexin-A level and lower total dendritic length and number in the dentate gyrus granule cells eight days after predator exposure. Phenotypically, this refers to a significant functional impairment that could affect reproduction and survival and thus fitness and population dynamics.

Large scale systemic control short-circuits pathogen transmission by interrupting the sand rat (Psammomys obesus)-to-sand fly (Phlebotomus papatasi) Leishmania major transmission cycle.

Systemic control uses the vertebrate hosts of zoonotic pathogens as "Trojan horses," killing blood-feeding female vectors and short-circuiting host-to-vector pathogen transmission. Previous studies focused only on the effect of systemic control on vector abundance at small spatial scales. None were conducted at a spatial scale relevant for vector control and none on the effect of systemic control on pathogen transmission rates. We tested the application of systemic control, using Fipronil-impregnated rodent baits, in reducing Leishmania major (Kinetoplastida: Trypanosomatidae; Yakimoff & Schokhor, 1914) infection levels within the vector, Phlebotomus papatasi (Diptera: Psychodidae; Scopoli, 1786) population, at the town-scale. We provided Fipronil-impregnated food-baits to all Psammomys obesus (Mammalia:Muridae; Cretzschmar, 1828), the main L. major reservoir, burrows along the southern perimeter of the town of Yeruham, Israel, and compared sand fly abundance and infection levels with a non-treated control area. We found a significant and substantial treatment effect on L. major infection levels in the female sand fly population. Sand fly abundance was not affected. Our results demonstrate, for the first time, the potential of systemic control in reducing pathogen transmission rates at a large, epidemiologically relevant, spatial scale.

Social behaviour and foraging success of little egrets (Egretta garzetta).

Intraspecific interactions among predators can change the game between a predator and its prey. Individuals of different size or sex can differ in their responses to conspecific competitors. We studied intraspecific interactions among pairs of little egrets (Egretta garzetta) while foraging on responsive prey (comet goldfish, Carassius auratus). Testing little egrets in an artificial patchy environment both singly and while engaged in social forging in pairs (male & female) at two prey densities, allowed us to explore individual differences in foraging success. We found sexual dimorphism with males being bigger and more aggressive than females. However, female foraging success was positively affected by the time they spent foraging with a conspecific male, suggesting they might be able to mitigate male aggressiveness with an indirect positive interaction. Despite the presence of direct interactions between individuals in the pair, egret foraging success was not affected by such interactions, nor by prey density. Our results demonstrate the importance of sex and an individual's ability to adjust its social behaviour based on the behaviour of others in this predator-prey foraging game.

The Potential of Systemic Control of Sand Flies Using Fipronil in the Novel Leishmania major (Kinetoplastida: Trypanosomatidae) Reservoirs Meriones tristrami (Rodentia: Muridea) and Meriones crassus (Rodentia: Muridea).

Leishmania major (Yakimoff & Schokhor, 1914), an important causative agent of Old World Cutaneous Leishmaniasis (CL), is transmitted by sand flies among a limited number of gerbilline reservoir-species. We can take advantage of this strong dependency to break the pathogen transmission cycle by using systemic insecticides that render the host toxic to the blood-feeding vector. We evaluated the potential of this approach with two novel reservoir species, incriminated for CL expansion in several sites in the Middle East. Specifically, we evaluated: 1) the residuality of the systemic insecticide fipronil in Meriones tristrami (Thomas, 1892) fed on fipronil-treated baits and 2) the treatments' adulticide effect on sand flies that blood fed on treated and untreated M. tristrami and M. crassus (Sundevall, 1842). We fed M. tristrami with food pellets containing 0.1 g/kg fipronil and used gas chromatograph-mass spectrometery analysis and bioassays to examine its residual toxicity to blood-feeding female sand flies. In M. tristrami, fipronil was rapidly metabolized to fipronil sulfone, found in the blood, urine, and feces for ≥31 d after fipronil admission. The survival of sand flies that blood fed on fipronil-treated M. tristrami and M. crassus was significantly reduced for at least 15 and 9 d respectively, after fipronil admission. These results hold promise for the potential contribution of systemic control approaches to CL integrated management strategies against novel CL (due to L. major) outbreaks in Israel and elsewhere.

Systemic Control of Cutaneous Leishmaniasis Sand-Fly Vectors: Fipronil-Treated Rodent Bait Is Effective in Reducing Phlebotomus papatasi (Diptera: Psychodidae) Female Emergence Rate From Rodent Burrows.

The strong dependency of some vectors on their host as a source of habitat can be viewed as a weak link in pathogen's transmission cycles using the vertebrate host as a 'Trojan horse' to deliver insecticides directly to the vector-host point of contact (hereafter 'systemic control'). This could, simultaneously, affect the survival of blood-feeding females and coprophagic larvae. Sand-flies, vectors of leishmaniasis worldwide, are often dependent on their bloodmeal host as a source of habitat and may therefore be good candidates for systemic control. In the present study, we field-tested this methodology by baiting Meriones crassus (Sundevall, 1842) (Rodentia:Muridea) with Fipronil-treated food pellets and evaluated its effect on reducing sand-fly emergence rate, in general, and of that of blood-fed females, in particular. We demonstrated 86% reduction in the abundance of female sand-flies that exit burrows of Fipronil-treated jirds, whereas male abundance was unaffected. Furthermore, whereas in control burrows 20% of the females were blood-fed, in treatment burrows no blood-fed females were detected. Sand-fly abundance outside the burrows was not affected by burrow treatment. This highlights the focal specificity of this method: affecting female sand-flies that feed on the reservoir host. This should result in the reduction of the pathogen transmission rate in the vicinity of the treated area by reducing the prevalence of leishmania-infected sand-fly females. These results hold promise for the potential of the systemic control approach in this system. Our next-step goal is to test this methodology at a large-scale cutaneous leishmaniasis control program.

Individual willingness to leave a safe refuge and the trade-off between food and safety: a test with social fish.

Refuges offer prey animals protection from predation, but increased time spent hiding can reduce foraging opportunities. Within social groups, individuals vary in their refuge use and willingness to forage in the presence of a predator. Here, we examine the relative foraging benefits and mortality costs associated with individual refuge use and foraging behaviour within groups of goldfish (Carassius auratus) under predation risk from an avian predator (little egret-Egretta garzetta). We assessed individual order of emergence from the refuge and participation over 15 group foraging outings, and assigned each fish a daily outing index score. The individual fish that emerged from the refuge earlier than the other group members and that participated in more outings received high outing index scores and consumed more food compared with fish that tended to emerge in posterior positions and participate in fewer outings. However, individual fish that attained high outing index scores suffered a higher risk of predation. Furthermore, the amount of time the egret spent at the pool affected group foraging behaviour: as predation risk increased, groups of fish consumed significantly less food. Our results exemplify the trade-off between foraging success and safety from predation that prey species regularly experience.

State-dependent foraging among social fish in a risky environment.

In the presence of a predator, foraging is a dangerous task. Social individuals can respond to risk by forming groups, benefiting from enhanced collective anti-predator behavior but suffering from increased conspicuousness to predators. Within groups, individuals exhibit variable foraging behavior. One important factor influencing risky foraging behaviour is current energetic state, and individuals must trade off food and safety by deciding when to leave a protected refuge in order to find food. We generated mixed groups of goldfish (Carassius auratus) containing equal numbers of underfed and well-fed individuals and examined individual refuge use and willingness to take risks venturing into risky foraging areas in the presence of an avian predator (little egret-Egretta garzetta). Underfed fish exhibited higher levels of risky behaviour by participating in more foraging outings and emerging from the refuge in frontal group positions, compared with well-fed individuals. As expected, underfed fish benefitted by consuming more food, but surprisingly did not experience higher rates of mortality. This may be due to the fact that the egret predator rarely captured the first fish to emerge from the refuge, preferentially attacked groups of three or more fish, and often captured fish in the chaotic period following a failed initial strike. We demonstrate how differences in energetic condition can influence risk-taking behaviours among social individuals that subsequently influence relative levels of foraging success and group fission-fusion dynamics. Moreover, our results illustrate the risk associated with foraging in larger groups.

Binary patch assessment by goldfish under safe and dangerous conditions.

Food resources can occur heterogeneously in space or time and differ in their abundances. A forager should be able to determine the value of a patch and choose optimally how to exploit it. However, patch choice and exploitation may be influenced by predation risk. Using the Giving up Densities (GUDs) technique, we evaluated goldfish patch assessment and choice in a binary patch choice experiment. We offered a pair of unequal food patches containing high and low food quantity. We quantified goldfish foraging behavior in the presence and absence of a predator. Goldfish groups equalized the GUDs in the two patches in safe environments but left higher GUDs in the rich patch under predation risk. The results suggest that goldfish can use both "patch assessment rule" and "fixed time rule" to exploit resource patches and trade off food and danger.

How does the presence of a conspecific individual change the behavioral game that a predator plays with its prey?

Behavioral games predators play among themselves may have profound effects on behavioral games predators play with their prey. We studied the behavioral game between predators and prey within the framework of social foraging among predators. We tested how conspecific interactions among predators (little egret) change the predator-prey behavioral game and foraging success. To do so, we examined foraging behavior of egrets alone and in pairs (male and female) in a specially designed aviary consisting of three equally spaced pools with identical initial prey (comet goldfish) densities. Each pool was comprised of a risky microhabitat, rich with food, and a safe microhabitat with no food, forcing the fish to trade off food and safety. When faced with two versus one egret, we found that fish significantly reduced activity in the risky habitat. Egrets in pairs suffered reduced foraging success (negative intraspecific density dependence) and responded to fish behavior and to their conspecific by changing their visiting regime at the different pools-having shorter, more frequent visits. The time egret spent on each visit allowed them to match their long-term capture success rate across the environment to their capture success rate in the pool, which satisfies one aspect of optimality. Overall, egrets in pairs allocated more time for foraging and changed their foraging tactics to focus more on fish under cover and fish 'peeping' out from their shelter. These results suggest that both prey and predator show behavioral flexibility and can adjust to changing conditions as needed in this foraging game.

A predator equalizes rate of capture of a schooling prey in a patchy environment.

Prey individuals are often distributed heterogeneously in the environment, and their abundances and relative availabilities vary among patches. A foraging predator should maximize energetic gains by selectively choosing patches with higher prey density. However, catching behaviorally responsive and group-forming prey in patchy environments can be a challenge for predators. First, they have to identify the profitable patches, and second, they must manage the prey's sophisticated anti-predator behavior. Thus, the forager and its prey have to continuously adjust their behavior to that of their opponent. Given these conditions, the foraging predator's behavior should be dynamic with time in terms of foraging effort and prey capture rates across different patches. Theoretically, the allocation of its time among patches of behaviorally responsive prey should be such that it equalizes its prey capture rates across patches through time. We tested this prediction in a model system containing a predator (little egret) and group-forming prey (common gold fish) in two sets of experiments in which (1) patches (pools) contained equal numbers of prey, or in which (2) patches contained unequal densities of prey. The egret equalized the prey capture rate through time in both equal and different density experiments.

Time budget, oxygen consumption and body mass responses to parasites in juvenile and adult wild rodents.

BACKGROUND: The study of changes in a host's energy allocation in response to parasites is crucial for understanding parasite impact on both individual- and population-level processes. Experimental studies have explored such responses mainly in a single subsample of hosts per study, primarily adult males, and have only assessed either the overall energy acquisition or expenditure, rather than their different components simultaneously, or the behavioral responses. Accordingly, two fundamental questions arise: why have multiple host strategies evolved to cope with increased energy expenditure? and, which factors determine this variation (e.g. host species, identity, age)? This study provides an important step towards addressing both questions by experimentally disentangling the short-term physiological and behavioral responses of juvenile and non-reproductive adult rodents to natural levels of flea infestation. These two cohorts represent extreme cases of the energy demand continuum, as the former, in contrast to the latter, is involved in growth--a highly energy-demanding process--and may not be able to operate far below its upper limit of energy expenditure, and thus should reduce its energy expenses upon the occurrence of extra demands (e.g. due to parasitic pressure). Accordingly, we hypothesized that the response to fleas is age-dependent and varies according to the age-specific energy requirements and constraints. METHODS: We monitored the behavior and physiology of juvenile and non-reproductive adult rodents before and after experimental flea infestation. First, we used a model selection approach to search for the factors that best explained the variability in the time budget, oxygen consumption, and body mass change in response to fleas. Then, using a path analysis approach, we quantified the different pathways connecting the important associations revealed at stage 1. RESULTS: Compared to their flea-free counterparts, flea-infested adults groomed longer and had a higher oxygen consumption rate, but did not lose body mass. Infested juveniles also groomed longer but grew slower and had a similar rate of oxygen consumption. CONCLUSIONS: Results suggest that both juvenile and adult rodents suffer from natural flea infestation levels. However, the comparison between the responses of juveniles and adults to experimental infestation, also suggests that juveniles may reallocate their energy expenditure from growth to maintenance, while non-reproductive adults increase their energy acquisition. Such age-dependent responses suggest that juveniles may be constrained by their higher need to rest for full functioning or by an upper limit in energy expenditure. Taken together, our study provides experimental evidence that hosts can compensate for the costs incurred by parasitism through physiological and behavioral plasticity, depending on their age, which probably determines their requirements and constraints. These compensatory responses may have important implications for the population dynamics of hosts and their parasites.

Effects of fish cues on mosquito larvae development.

We investigated the effects of predator-released kairomones on life history traits of larval Culex pipiens (Linnaeus). We compared the development time and survival of sibling larvae, reared in either water conditioned by the presence of Gambusia affinis (Baird and Girard) or fishless control-water. Our results indicate that larvae developing in fish-conditioned water (FCW) pupated faster than larvae in fishless-control water. The effect of FCW on larval survival was evident only in females. Surprisingly, FCW increased female survival. In both development-time and survival, boiling the water eliminated the FCW effect, supporting our hypothesis that fish conditioning is based on kairomones. Accelerated metamorphosis in response to predator released kairomones, evident in our results, is a rarely described phenomenon. Intuitively, when exposed to predator associated signals, aquatic larvae should metamorphose earlier to escape the higher risk of predation. However, theoretical models predict this outcome only under specific conditions. Indeed, longer - rather than shorter - time to metamorphosis is usually observed in response to predation risk. We argue that the response of larval mosquitoes to predation risk is context-dependent. Shortening larval development time may not be an exceptional response, but rather represents a part of a response spectrum that depends on the level of predation risk and resource abundance.

Jack of all trades, master of all: a positive association between habitat niche breadth and foraging performance in pit-building antlion larvae.

Species utilizing a wide range of resources are intuitively expected to be less efficient in exploiting each resource type compared to species which have developed an optimal phenotype for utilizing only one or a few resources. We report here the results of an empirical study whose aim was to test for a negative association between habitat niche breadth and foraging performance. As a model system to address this question, we used two highly abundant species of pit-building antlions varying in their habitat niche breadth: the habitat generalist Myrmeleon hyalinus, which inhabits a variety of soil types but occurs mainly in sandy soils, and the habitat specialist Cueta lineosa, which is restricted to light soils such as loess. Both species were able to discriminate between the two soils, with each showing a distinct and higher preference to the soil type providing higher prey capture success and characterizing its primary habitat-of-origin. As expected, only small differences in the foraging performances of the habitat generalist were evident between the two soils, while the performance of the habitat specialist was markedly reduced in the alternative sandy soil. Remarkably, in both soil types, the habitat generalist constructed pits and responded to prey faster than the habitat specialist, at least under the temperature range of this study. Furthermore, prey capture success of the habitat generalist was higher than that of the habitat specialist irrespective of the soil type or prey ant species encountered, implying a positive association between habitat niche-breadth and foraging performance. Alternatively, C. lineosa specialization to light soils does not necessarily confer upon its superiority in utilizing such habitats. We thus suggest that habitat specialization in C. lineosa is either an evolutionary dead-end, or, more likely, that this species' superiority in light soils can only be evident when considering additional niche axes.

Driven to distraction: detecting the hidden costs of flea parasitism through foraging behaviour in gerbils.

Gerbilline rodents such as Allenby's gerbils (Gerbillus andersoni allenbyi), when parasitized by fleas such as Synosternus cleopatrae pyramidis, devote long hours of grooming to remove the ectoparasites. Yet no detrimental energetic or immunological effects of the ectoparasites have been found in adult Allenby's gerbil. Why should gerbils go to such trouble? We tested for the various ways that fleas can negatively affect gerbils by manipulating flea infestation on gerbils and the presence of a fox. We demonstrate that gerbils responded to fleas by leaving resource patches at higher giving-up densities. Furthermore, they stayed in those resource patches less time and left them at higher quitting harvest rates so long as a fox was also present. When flea-ridden, gerbils also abandoned using vigilance to manage risk and relied mainly on time allocation. Thus, having fleas imposed a foraging cost similar in nature to that arising from the risk of predation from foxes and may be even larger in magnitude. More than that, the presence of fleas acted as a magnifier of foraging costs, especially those arising from the risk of predation. The fleas reduced the gerbils' foraging aptitude and altered how they went about managing risk of predation. We hypothesize that fleas reduce the attention that gerbils otherwise have for foraging and predator detection. We suggest that this is the major cost of ectoparasitism.

Conservation genetics of a rare Gerbil species: a comparison of the population genetic structures and demographic histories of the locally rare Pygmy Gerbil and the common Anderson's Gerbil.

BACKGROUND: One of the major challenges in evolutionary biology is identifying rare species and devising management plans to protect them while also sustaining their genetic diversity. However, in attempting a broad understanding of rarity, single-species studies provide limited insights because they do not reveal whether the factors that affect rare species differ from those that affect more common species. To illustrate this important concept and to arrive at a better understanding of the form of rarity characterizing the rare Gerbillus henleyi, we explored its population genetic structure alongside that of the locally common Gerbillus andersoni allenbyi. We trapped gerbils in several locations in Israel's western and inner Negev sand dunes. We then extracted DNA from ear samples, and amplified two mitochondrial sequences: the control region (CR) and the cytochrome oxidase 2 gene (CO2). RESULTS: Nucleotide diversity was low for all sequences, especially for the CR of G. a. allenbyi, which showed no diversity. We could not detect any significant population genetic structure in G. henleyi. In contrast, G. a. allenbyi's CO2 sequence showed significant population genetic structure. Pairwise PhiPT comparisons showed low values for G. henleyi but high values for G. a. allenbyi. Analysis of the species' demographic history indicated that G. henleyi's population size has not changed recently, and is under the influence of an ongoing bottleneck. The same analysis for G. a. allenbyi showed that this species has undergone a recent population expansion. CONCLUSIONS: Comparing the two species, the populations of G. a. allenbyi are more isolated from each other, likely due to the high habitat specificity characterizing this species. The bottleneck pattern found in G. henleyi may be the result of competition with larger gerbil species. This result, together with the broad habitat use and high turnover rate characterizing G. henleyi, may explain the low level of differentiation among its populations. The evidence for a recent population expansion of G. a. allenbyi fits well with known geomorphological data about the formation of the Negev sand dunes and paleontological data about this species' expansion throughout the Levant. In conclusion, we suggest that adopting a comparative approach as presented here can markedly improve our understanding of the causes and effects of rarity, which in turn can allow us to better protect biodiversity patterns.

Foraging behavior of an urban bird species: molt gaps, distance to shelter, and predation risk.

Predation cost (Pc) is often regarded as a pivotal component determining foraging behavior. We hypothesized that variations in two of its major constituents, predation risk (mu) and the marginal value of energy ([see text for symbol]Fs/ [see text for symbol]e, where Fs is the survivor's fitness and e represents the amount of acquired energy), will translate into variations in patch use behavior of ground-foraging birds. We studied patch use behavior of House Sparrows (Passer domesticus), as affected by the proximity to shelter, in two large outdoor aviaries. Proximity to shelter should affect mu. We manipulated the birds' flight performance by clipping primary flight feathers from their wings to increase mu, but the clipping may also increase [see text for symbol]Fs/ [see text for symbol]e. To help distinguish between the birds' response to these confounding effects, we further augmented food in the aviaries to reduce [see text for symbol]Fs/ [see text for symbol]e. Patch use, as measured by giving-up densities (GUD, the amount of food left behind in a resource patch following exploitation) was affected by distance from shelter only slightly and mainly when the birds were feather-clipped and food was not augmented. Food augmentation had a homogenizing effect on foraging costs by increasing GUDs and washing out the effects of distance and feather clipping. We argue that mu increases with distance from shelter but that, for the highly urban House Sparrow, this increase is only slight. Feather clipping then increased mu further to the point at which patch use discernibly decreased with distance from shelter. Our experimental manipulation of feather clipping also acted to increase [see text for symbol]Fs/ [see text for symbol]e and resulted in an overall lowering of GUDs. The seed augmentation counteracted the effect of feather clipping on [see text for symbol]Fs/ [see text for symbol]e, allowing the birds to reduce their foraging efforts and washing out the qualitative effect of mu with respect to distance from shelter.

Ecological trap for desert lizards caused by anthropogenic changes in habitat structure that favor predator activity.

Anthropogenic habitat perturbation is a major cause of population decline. A standard practice managers use to protect populations is to leave portions of natural habitat intact. We describe a case study in which, despite the use of this practice, the critically endangered lizard Acanthodactylus beershebensis was locally extirpated from both manipulated and natural patches within a mosaic landscape of an afforestation project. We hypothesized that increased structural complexity in planted patches favors avian predator activity and makes these patches less suitable for lizards due to a heightened risk of predation. Spatial rarity of natural perches (e.g., trees) in arid scrublands may hinder the ability of desert lizards to associate perches with low-quality habitat, turning planted patches into ecological traps for such species. We erected artificial trees in a structurally simple arid habitat (similar to the way trees were planted in the afforestation project) and compared lizard population dynamics in plots with these structures and without. Survival of lizards in the plots with artificial trees was lower than survival in plots without artificial trees. Hatchlings dispersed into plots with artificial trees in a manner that indicated they perceived the quality of these plots as similar to the surrounding, unmanipulated landscape. Our results showed that local anthropogenic changes in habitat structure that seem relatively harmless may have a considerable negative effect beyond the immediate area of the perturbation because the disturbed habitat may become an ecological trap.

Effects of food abundance, age, and flea infestation on the body condition and immunological variables of a rodent host, and their consequences for flea survival.

Temporal variation in body condition and immunological variables of animals that harbor parasites may explain patterns of variation in infestation, as well as parasite impact on the host. We emulated such variability in Sundevall's jirds by manipulating food availability and flea infestation in juveniles and adults and examining how these changes affect survival of fleas on their hosts. Body condition of food-restricted jirds deteriorated, but there was no change in their immunological variables. Adult jirds were in better body condition and had higher immunocompetence than juveniles, however there were no significant effects of flea infestation on any of the variables examined. The main effects of flea infestation were a decrease in the response to phytohaemagglutinin injection, and an increase in the negative effects of food restriction on body mass. Flea survival was higher on juveniles, but fleas did not respond to temporal variability in body condition and immunocompetence of the jirds. We concluded that changes in body condition and immune responses due to growth or variability in food abundance are more important than changes caused by the fleas themselves. Flea infestation is more detrimental to jirds when they are not able to compensate for mass loss through increased food consumption.

Ultimate mechanisms of age-biased flea parasitism.

Mechanisms that cause nonrandom patterns of parasite distribution among host individuals may influence the population and evolutionary dynamics of both parasites and hosts, but are still poorly understood. We studied whether survival, reproduction, and behavioral responses of fleas (Xenopsylla conformis) changed with the age of their rodent hosts (Meriones crassus), experimentally disentangling two possible mechanisms: (a) differential survival and/or fitness reward of parasites due to host age, and (b) active parasite choice of a host of a particular age. To explore the first mechanism, we raised fleas on rodents of two age groups and assessed flea survival as well as the quantity and quality of their offspring. To explore the second mechanism, three groups of fleas that differed in their previous feeding experience (no experience, experience on juvenile or experience on adult rodents) were given an opportunity to choose between juvenile and adult rodents in a Y-maze. Fleas raised on juvenile rodents had higher survival and had more offspring that emerged earlier than fleas raised on adults. However, fleas did not show any innate preference for juvenile rodents, nor were they able to learn to choose them. In contrast to our predictions, based on a single previous exposure, fleas learned to choose adult rodents. The results suggest that two mechanisms-differential survival and fitness reward of fleas, and associative learning by them-affect patterns of flea distribution between juvenile and adult rodents. The former increases whereas the latter reduces flea densities on juvenile rodents. The ability of fleas to learn to choose adult but not juvenile hosts may be due to: (a) a stronger stimulus from adults, (b) a higher profitability of adults in terms of predictability and abundance, or (c) the evolutionary importance of recognizing adult but not juvenile hosts as representatives of the species.