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.     

Measuring the benefit of habitat selection

We used a behavioral bioassay to estimate the advantages thattwo species of gerbils (Gerbillus allenbyi and G. pyramidum)experienced by preferring a semistabilized dune habitat overa stabilized sand habitat. We used the magnitude of foragingeffort by the gerbils to signal the difference between thetwo habitats. When they were foraging as much in stabilizedsand as in semistabilized dune, we inferred that these habitatswere providing equivalent rewards. We performed a series ofexperiments in two 1-ha field enclosures, each containing similarproportions of stabilized sand and semistabilized dune. Eachenclosure contained a population of only one of the species.By varying the amount of seeds added (either 0.5, 1, 2, or 3g of seeds in 18 seed trays) to each habitat and monitoringthe behavior of the gerbils, we were able to fit a curve thatreflected the change in habitat preference as a function ofseed addition rate. We were also able to show how much seedaddition had to be added to bring the two habitats into equaluse. Each species required only 13 g/ha/night to entirely offsetthe advantage of the semistabilized dune.     

Can interaction coefficients be determined from cencus data?

Summary The method of estimating interactions proposed independently by Pimm and Schoener is studied using field data from the community of rodents which lives in the arid, rocky habitats of Israel. One important problem the method addresses is how to remove the effects of habitat heterogeneity on the estimate. We tried six different variations of the analysis scheme outlined by Crowell and Pimm, and found their results qualitatively inconsistent. This was especially true when we compared the results produced from separate habitat variables with those produced from the principal components of the habitat variation.Another problem, this one not previously addressed, is great variation in the average abundance of the different species. We discovered that the ratio of the average abundances of two species is the best predictor of the value of their coefficients of interaction. Common species appear to have weak influence on rare ones; rare ones appear to have strong influence on common ones. The statistical mechanism which produces this relationship is clear, indicating that the relationship is an artifact.     

Two gerbils of the Negev: A long-term investigation of optimal habitat selection and its consequences

Optimal foraging theory has entered a new phase. It is not so much tested as used. It helps behavioural ecologists discover the nature of the information in an animals brain. It helps population ecologists reveal coefficients of interaction and their patterns of density-dependent variation. And it helps community ecologists examine niche relationships. In our studies on two species of Negev desert gerbil, we have taken advantage of the second and third of these functions. Both these gerbils prefer semi-stabilized dune habitat, and both altered their selective use of this habitat and stabilized sand according to experimental changes we made in their populations. Their changes in selectivity agree with a type of optimal foraging theory called isoleg theory. Isoleg theories provide examples of dipswitch theories – bundles of articulated qualitative predictions – that are easier to falsify than single qualitative predictions. By linking behaviour to population dynamics through isoleg theory, we were able to use the behaviour of the gerbils to reveal the shapes of their competitive isoclines. These have the peculiar non-linear shapes predicted by optimal foraging theory. Finally, when owl predation threatens, the behaviour of Gerbillus allenbyi reveals the shape of their victim isocline. As has long been predicted by predation theory and laboratory experiments, it is unimodal.     

Inter-specific competitors reduce inter-gender competition in Negev Desert gerbils

We examined gender-dependent competitive interactions between two nocturnal desert gerbil species, Gerbillus andersoni allenbyi and G. pyramidum, by a field manipulation experiment. The study was done in two 1-ha enclosed plots and included allopatric (only G. a. allenbyi) and sympatric (both species together) treatments. Seed trays and thermal imaging cameras were used to observe the gerbils foraging activities and aggressive interactions. We found that the negative effect of the competitively dominant species, G. pyramidum, on time spent in seed trays, and ability to control these artificial food patches, was stronger on male than on female G. a. allenbyi. Consequently, the aggression of male G. a. allenbyi towards female G. a. allenbyi was markedly reduced, indicating that the dominant species mediated competition between the genders of the subordinate species. Furthermore, this interference-mediated indirect effect was associated with a decrease in the body mass of male G. a. allenbyi and an increase in the survival of female G. a. allenbyi. We suggest that both the reduction in intra-specific aggression and the positive effect on female survival can potentially stabilize competitive interactions and promote coexistence in this small mammal community.     

Benefits, Costs and Constraints of Anti-Parasitic Grooming in Adult and Juvenile Rodents

Grooming behaviour plays various roles in the health care, reproduction, and social life of an individual vertebrate. However, the reasons for the variability in time spent grooming amongst species, populations and individuals are not fully understood. We tested the hypothesis that the main role of grooming is ectoparasite removal and thus that time spent grooming by an animal reflects the costs of parasite infestation offset against the costs of grooming. The test was conducted on a rodent, Meriones crassus, that is parasitised by a flea, Xenopsylla conformis. We monitored behaviour of juvenile and adult rodents before and after flea infestation and quantified the probability of mortality of fleas with respect to the time spent grooming in adults compared with juvenile rodents. We predicted that: (1) increased costs of flea infestation (e.g. in parasitised as opposed to flea‐free rodents and in juveniles as opposed to adults) increases time spent grooming and (2) mortality probability per flea increases with increasing time spent grooming and is higher for fleas on juveniles than for fleas on adult rodents. We were interested to discover at the expense of which activity grooming is increased. Our findings established that the major role of grooming is in flea removal, as exposure to fleas evoked grooming activity in all rodents and grooming activity explained 57–70% of the variation in flea mortality. Furthermore, we showed that the rise in grooming activity was at the expense of resting. However, we found only partial support for the predicted increase in grooming time with increasing costs of flea infestation. Flea infestation did indeed increase the time spent grooming by rodents. Nevertheless, juvenile rodents who incur higher costs of flea infestation spent less time grooming than adults and sustained similar flea densities, suggesting that these hosts are constrained by some other factors, such as feeding time.