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.

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.

Decoupling fragmentation from habitat loss for spiders in patchy agricultural landscapes.

Habitat loss reduces species diversity, but the effect of habitat fragmentation on number of species is less clear because fragmentation generally accompanies loss of habitat. We compared four methods that aim to decouple the effects of fragmentation from the effects of habitat loss. Two methods are based on species-area relations, one on Fisher's alpha index of diversity, and one on plots of cumulative number of species detected against cumulative area sampled. We used these methods to analyze the species diversity of spiders in 2, 3.2 × 4 km agricultural landscapes in Southern Judea Lowlands, Israel. Spider diversity increased as fragmentation increased with all four methods, probably not because of the additive within-patch processes, such as edge effect and heterogeneity. The positive relation between fragmentation and species diversity might reflect that most species can disperse through the fields during the wheat-growing season. We suggest that if a given area was designated for the conservation of spiders in Southern Judea Lowlands, Israel, a set of several small patches may maximize species diversity over time.

Habitat fragmentation may not matter to species diversity.

Conservation biologists worry that fragmenting a bloc of natural habitat might reduce its species diversity. However, they also recognize the difficulty and importance of isolating the effect of fragmentation from that of simple loss of area. Using two different methods (species-area curve and Fisher's alpha index of diversity) to analyse the species diversities of plants, tenebrionid beetles and carabid beetles in a highly fragmented Mediterranean scrub landscape, we decoupled the effect of degree of fragmentation from that of area loss. In this system, fragmentation by itself seems not to have influenced the number of species. Our results, obtained at the scale of hectares, agree with similar results at island and continent scales.

Clump size selection: A field test with two species of Dipodomys.

Seed distribution (clump size) selection has been proposed as a possible mechanism of resource subdivision for competing heteromyid rodent species. To test this hypothesis, field experiments were conducted over two years during both richer and sparser seasons of the year. None of the predictions derived from the hypothesis were supported by our results. Though some selectivity was displayed by both Dipodomys spectabilis and D. merriami, the patterns of selectivity did not match the expected patterns. Our results further indicate that clump selection may be influenced by variables other than the density of seeds within a clump. These results have led us to conclude that clump size selection is unlikely to play a role in the coexistence of different species of the genus Dipodomys.

Perturbation analysis of competition and overlap in habitat utilization between Dipodomys ordii and Dipodomys merriami.

The populations of two coexisting species of Dipodomys (Heteromyidae, Rodentia) were manipulated on 10, large, unenclosed, trapping grids. These manipulations revealed that, although many kangaroo rats are established residents in an area, a large number are transient individuals who quickly occupy vacated habitats. On plots from which residents had been removed, transients settled at rates of up to 5% of carrying capacity per day. These immigrants were invariably of the same species that was removed, indicating a strong element of intraspecific competition with little or no evidence of competition between the species.Trapping records suggest that these species avoid competition through habitat selection. Dipodomys ordii prefer a grassier habitat, and D. merriami a habitat dominated by creosote bush. Apparent overlap in their utilization of habitats, based on sites of capture, predicts competition coefficients to be higher than those permitted by the theory of limiting similarity and much higher than those actually shown by the perturbation experiments.This study demonstrates the dangers of estimating alpha without experimentation. This is especially true in cases where habitat selection may be important, since organisms may travel in habitats without collecting resources therein. Our results are discussed in light of a theory which examines the optimal (rather than tolerable) amount of overlap in habitat utilization between two potential competitors in a mixed habitat. This theory predicts that the pressure of natural selection should eliminate the interspecific competition entirely.However, the conclusion that the interspecific competitive alpha is zero does not lead to the conclusion that interspecific competition is unimportant in the system. Instead, if our interpretation is correct, such competition has molded the system, and were there not a continual threat of interspecific competition, the habitat specializations would soon disappear.

Microhabitat selection in two species of heteromyid rodents.

An experiment was conducted to determine the microhabitat preferences of two heteromyid rodents, Dipodomys ordi and Perognathus flavus. This experiment used marked seeds and the atomic absorption spectrophotometer in order to study the environment as a mosiac of microhabitats. The results of our analysis indicate that these two heteromyids are microhabitat selectors. The preferences of the rodents are D. ordi: grass habitat 0.0%, near grass habitat 22.5%, open habitat 77.4% and P. flavus: grass habitat 46.2%, near grass habitat 32.2%, open habitat 21.4%. The overlap between the two species is only 0.43.

Can interaction coefficients be determined from cencus data?

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.