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Allee effects, extinctions, and chaotic transients in simple population models.

Discrete time single species models with overcompensating density dependence and an Allee effect due to predator satiation and mating limitation are investigated. The models exhibit four behaviors: persistence for all initial population densities, bistability in which a population persists for intermediate initial densities and otherwise goes extinct, extinction for all initial densities, and essential extinction in which "almost every" initial density leads to extinction. For fast-growing populations, these models show populations can persist at high levels of predation even though lower levels of predation lead to essential extinction. Alternatively, increasing the predator's handling time, the population's carrying capacity, or the likelihood of mating success may lead to essential extinction. In each of these cases, the mechanism behind these disappearances are chaotic dynamics driving populations below a critical threshold determined by the Allee effect. These disappearances are proceeded by chaotic transients that are proven to be approximately exponentially distributed in length and highly sensitive to initial population densities.

Alleles↗

Exploitation of an ancient escape circuit by an avian predator: prey sensitivity to model predator display in the field.

Certain insectivorous birds, such as the painted redstart (Myioborus pictus), undertake flush pursuit--a characteristic display that elicits an escape reaction by an insect, which the bird then chases in the air and eats. This account describes experiments showing that flush pursuit uses visual displays, which are likely to exploit an ancient neural circuit in dipteran insects, the visual systems of which are well documented as detecting looming stimuli and triggering an escape responses. Using models that decompose components of the redstart display, specific elements of the display were analyzed for their contribution in triggering visually induced escape behavior by dipterous insects. Elements tested were pivoting body movements, patterning on the spread tail and wings, and visual contrast of model redstarts against pale and dark backgrounds. We show that contrasting patterns within the plumage are crucial to foraging success, as is contrast of the bird against a background. Visual motion also significantly contributes to the successful flushing. In contrast, unpatterned models and patterned models that do not contrast with the background are less successful in eliciting escape responses of flies. Natural visual stimuli provided by Myioborus pictus are similar to those known to trigger looming and time-to-collision neurons in the escape circuits of flies and other insects, such as orthopterans. We propose that the tuning properties of these neural pathways might have contributed to the evolution of foraging displays in flush-pursuing birds.

Animals↗

The structural stability of a three-species food chain model.

A three-species food-chain model which was previously shown to exhibit chaotic dynamics was revisited. By exploring the sensitivity of that result this study found that complex behavior depended on the functional form chosen to model the interaction between the two highest species in the food chain. Two separate scenarios were explored: the gradual addition of refugia modeling the escape from predation at low prey densities; and the gradual addition of predator interference modeling territorial behavior. The addition of even a small amount of refugia provided a stabilizing influence as the chaotic dynamics collapsed to stable limit cycles. The results of adding interference to the model were more complex. Although the numerical simulations indicated that a low level of interference provided a stabilizing influence, the analytical results suggest that complex dynamics are possible for a range of parameter values that are biologically relevant. The sensitivity of the stability profile to functional changes in the model suggests two important ecological motivations for structural stability analysis. First, in ecological systems, environmental fluctuations cause continuous changes in the functional relationships between and within species, resulting in potential changes in the complexity of the dynamics over time. Second, slight changes in ecological structure may cause significant bifurcations; however, most ecological data are inadequate to distinguish such phenomena.

Animals↗

Age differences in the response of California ground Squirrels (Spermophilus beecheyi) to avian and mammalian predators.

The antipredator behavior of juvenile and adult California ground squirrels (Spermophilus beecheyi) was videotaped in Experiment 1 to measure the effects of age on assessment of a briefly presented live dog and a model red-tailed hawk (Buteo jamaicensis) in simulated flight. Adult squirrels treated the hawk as more dangerous than the dog, whereas juvenile squirrels showed less differentiation of the predator types. Juvenile squirrels also perceived the dog as a more immediate danger than adult squirrels did. For Experiment 2, the red-tailed hawk model was compared with models of a nonthreatening turkey vulture (Cathartes aura) and crow (Corvus brachyrhynchos). Neither age class differentiated the avian models; however, the adult squirrels treated these birds as more threatening than the juvenile squirrels did. Both studies suggest that learning may contribute to predator assessment.

Aging↗

Parametric analysis of a predator-prey system stabilized by a top predator.

We present a complete parametric analysis of a predator-prey system influenced by a top predator. We study ecosystems with abundant nutrient supply for the prey where the prey multiplication can be considered as proportional to its density. The main questions we examine are the following: (1) Can the top predator stabilize such a system at low densities of prey? (2) What possible dynamic behaviors can occur? (3) Under which conditions can the top predation result in the system stabilization? We use a system of two nonlinear ordinary differential equations with the density of the top predator as a parameter. The model is investigated with methods of qualitative theory of ODEs and the theory of bifurcations. The existence of 12 qualitatively different types of dynamics and complex structure of the parametric space are demonstrated. Our studies of phase portraits and parametric diagrams show that a top predator can be an important factor leading to stabilization of the predator-prey system with abundant nutrient supply. Although the model here is applied to the plankton communities with fish (or carnivorous zooplankton) as the top trophic level, the general form of the equations allows applications of our results to other ecological systems.

Algorithms↗

Food chain dynamics in the chemostat.

The asymptotic behavior of a tri-trophic food chain model in the chemostat is studied. The Monod-Herbert growth model is used for all trophic levels. The analysis is carried out numerically, by finding both local and global bifurcations of equilibria and of limit cycles with respect to two chemostat control parameters: the dilution rate of the chemostat and the concentration of input substrate. It is shown that the bifurcation structure of the food chain model has much in common with the bifurcation structure of a one-dimensional map with two turning points. This map is used to explain how attractors are created and destroyed under variation of the bifurcation parameters. It is shown that low as well as high concentration of input substrate can lead to extinction of the highest trophic level.

Animals↗

Daphnia growth is hindered by chemical information on predation risk at high but not at low food levels.

Daphnia hyalina x galeata (Dhg) and D. pulicaria (Dp) are ready to pay greater costs in terms of predation risk avoidance at high rather than at low food levels. Such costs are easier to assess in Daphnia than in large long-lived and difficult-to-handle herbivores, since they can be precisely determined in a few-day experiment as the reduced growth (P=A-R) resulting from diminished assimilation (A) and/or increased respiration (R). In experiments with Daphnia grown for six days from the neonate to the first clutch of eggs, which were given different levels of algal food (Scenedesmus at concentrations from 0.05 to 1.60 mg C l(-1)), individual growth was lower in the presence of fish kairomone (chemical information on fish predation; present at a concentration that induces antipredator defensive behavior and life histories) than in the absence of kairomone (control). The difference from the control was negligible at the lowest food levels, and gradually increased with increasing food concentration. At a food concentration of 1.6 mg C l(-1), growth was reduced by 9-32 and 0-8% in Dhg and Dp, respectively, compared to the controls. A similar reduction was observed in the body length of six-day-old animals (Dhg 6-19%, Dp 0-14%), but not in the first clutch reproductive effort (clutch volume). Daphnia had a greater number of eggs per clutch in the presence of the kairomone, but smaller eggs, so that the total volume of eggs in a clutch was the same with and without kairomone. The amplification of the effect of the kairomone due to high food levels was weaker in Dp, a species that rarely coexists with planktivorous fish in natural habitats.

Animals↗

Trichobothrial mediation of an aquatic escape response: directional jumps by the fishing spider, Dolomedes triton, foil frog attacks.

Fishing spiders (Pisauridae) frequent the surfaces of ponds and streams and thereby expose themselves to predation by a variety of aquatic and semi-aquatic vertebrates. To assess the possibility that the impressive jumps of fishing spiders from the water surface function in evading attacks by frogs, attacks by bullfrogs (Rana catesbiana) and green frogs (R. clamitans) on Dolomedes triton were studied. Both the attack dynamics of the frogs and the evasive behaviors of the spiders were recorded at 250 frames per second. A freeze-dried bullfrog, propelled toward spiders with acceleration, posture, and position that approximated the natural attack posture and dynamics, was used to assess the spiders' behavior. Qualitatively, the spiders responded to these mock-attacks just as they had to attacks by live frogs: jumping (N=29 jumps, 56.9% of instances), rearing the legs nearest the attacking frog (N=15, 29.4%), or showing no visible response (N=7, 13.7%). Spiders that jumped always did so away (in the vertical plane) from the attack (mean =137 degrees vs. vertical at 90 degrees or horizontally toward the frog at 0 degrees ). The involvement of the trichobothria (leg hairs sensitive to air movements), and the eyes as sensory mediators of the evasion response was assessed. Spiders with deactivated trichobothria were significantly impaired relative to intact and sham-deactivated spiders, and relative to spiders in total darkness. Thus, functional trichobothria, unlike the eyes, are both necessary and sufficient mediators of the evasion response. Measurements of air flow during frog attacks suggest that an exponential rise in flow velocity is the airborne signature of an attack.

Animals↗

Bistability and an Allee effect as emergent consequences of stage-specific predation.

The Allee effect, a reduction of individual fitness at low population density that can lead to sudden and unannounced extinctions, has been shown to come about through a number of mechanisms, usually associated with group behavior or mate search. Recent papers show that it may arise through size-selective predation, without explicit assumptions relating individual fitness to population density. It arises from the shift that a predator induces in the population stage distribution of its prey. We study the parameter conditions that lead to such an emergent Allee effect. The emergent Allee effect occurs under fairly broad conditions. We show that stage-specific predation can also induce bistability between alternative states where both prey and predator are present. A perturbation analysis on the equilibria shows that all equilibria are highly robust to changes in predator density. Our work shows that when size-specific interactions are taken into account, bistabilities and catastrophic collapses are possible even in purely exploitative food webs, which has substantial implications for questions related to food web theory and conservation issues.

Animals↗

Field verification of predator attraction to minnow alarm substance.

Fishes such as minnows in the superorder Ostariophysi possess specialized alarm substance cells (ASC) that contain an alarm cue. Alarm substance can only be released by damage to the epidermis; thus, the release of alarm substance is a reliable indicator of predation risk. When nearby minnows detect the cue, they adopt a range of antipredator behaviors that reduce their probability of predation. Predator-predator interactions afford prey an opportunity to escape and, thus, a fitness benefit that maintains alarm substance calls over evolutionary time. Here, we present data from a simple field experiment verifying that nearby predators are attracted to minnow alarm substance because it signals an opportunity to pirate a meal. Fishing lures were baited with sponge blocks scented with either (1) water (control for sponge odor and appearance), (2) skin extract from non-ostariophysan convict cichlids (superorder Acanthopterygii, Archocentrus "Cichlasoma" nigrofasciatus) to control for general injury-released cues from fish, or (3) skin extract from fathead minnows (superorder Ostariophysi, Pimephales promelas). Predator strike frequency on each sponge type was 1, 1, and 7 for water, cichlid, and minnow cues, respectively. These data provide the first field test using fish predators of the predator-attraction hypothesis for the evolution of Ostariophysan alarm substance cells.

Adaptation, Physiological↗

Ecological consequences of chemically mediated prey perception.

To locate food, mobile consumers in aquatic habitats perceive and move towards sources of attractive chemicals. There has been much progress in understanding how consumers use chemicals to identify and locate prey despite the elusive identity of odor signals and the complex effects of turbulence on chemical dispersion. This review highlights how integrative studies on behavior, fluid physics, and chemical isolation can be fundamental in elucidating mechanisms that regulate species composition and distribution. We suggest three areas where further research may yield important ecological insights. First, although basic aspects of stimulatory molecules are known, our understanding of how consumers identify prey from a distance remains poor, and the lack of studies examining the influence of distance perception on food preference may result in inaccurate estimation of foraging behavior in the field. Second, the ability of many animals to find prey is greatest in unidirectional, low turbulence flow environments, although recent evidence indicates a trade-off in movement speed versus tracking ability in turbulent conditions. This suggests that predator foraging mode may affect competitive interactions among consumers, and that turbulence provides a hydrodynamic refuge in space or time, leading to particular associations between predator success, prey distributions, and flow. Third, studies have been biased towards examining predator tracking. Current data suggest a variety of mechanisms prey may use to disguise their presence and avoid predation; these mechanisms also may produce associations between prey distributions and flow environments. These examples of how chemical attraction may mediate interactions between consumers and their resources suggest that the ecology of chemically mediated prey perception may be as fundamental to the organization of aquatic communities as the ecology of chemical deterrence.

Animals↗

A decade of discoveries in veterinary protozoology changes our concept of "subclinical" toxoplasmosis.

One of the most compelling topics to emerge from the last decade of veterinary protozoology is disease caused by a zoonotic pathogen, Toxoplasma gondii, in otherwise healthy people. These findings may catch the health professions by surprise, because veterinary and medical courses and textbooks typically emphasize that T. gondii infections are subclinical, unless acquired in utero or the patient has a serious immunosuppressive condition. Nevertheless, numerous reports in the last decade associate toxoplasmosis with lymphadenopathy, fever, weakness and debilitation, ophthalmitis, and severe multisystemic infections in people who do not have immunosuppressive conditions. Toxoplasmosis in rodents causes altered behavior, and similar mental aberrations are coming to light in humans; recent studies associate T. gondii infection with personality shifts and increased likelihood of reduced intelligence or schizophrenia. These conditions reduce the quality of life of individuals, and may exact a significant economic burden upon society. Of course, toxoplasmosis continues to cause serious conditions in AIDS patients and congenitally infected people, as well as abortions and encephalitis in domestic and wild animals. Environmental contamination is heavy enough to extend into marine wildlife. It is time for the health professions to amend teaching curricula regarding T. gondii. Veterinary parasitologists should lead the way in developing methods to reduce the prevalence of T. gondii in food animals. Public health policies should prohibit the practice of allowing pet cats to roam. Organizations and individuals that feed feral cats are unwittingly contributing to the dissemination of T. gondii, by sustaining artificially dense populations of a definitive host of this protozoal parasite.

Animals↗

Protozoan predation, diversifying selection, and the evolution of antigenic diversity in Salmonella.

Extensive population-level genetic variability at the Salmonella rfb locus, which encodes enzymes responsible for synthesis of the O-antigen polysaccharide, is thought to have arisen through frequency-dependent selection (FDS) by means of exposure of this pathogen to host immune systems. The FDS hypothesis works well for pathogens such as Haemophilus influenzae and Neisseria meningitis, which alter the composition of their O-antigens during the course of bloodborne infections. In contrast, Salmonella remains resident in epithelial cells or macrophages during infection and does not have phase variability in its O-antigen. More importantly, Salmonella shows host-serovar specificity, whereby strains bearing certain O-antigens cause disease primarily in specific hosts; this behavior is inconsistent with FDS providing selection for the origin or maintenance of extensive polymorphism at the rfb locus. Alternatively, selective pressure may originate from the host intestinal environment itself, wherein diversifying selection mediated by protozoan predation allows for the continued existence of Salmonella able to avoid consumption by host-specific protozoa. This selective pressure would result in high population-level diversity at the Salmonella rfb locus without phase variation. We show here that intestinal protozoa recognize antigenically diverse Salmonella with different efficiencies and demonstrate that differences solely in the O-antigen are sufficient to allow for prey discrimination. Combined with observations of the differential distributions of both serotypes of bacterial species and their protozoan predators among environments, our data provides a framework for the evolution of high genetic diversity at the rfb locus and host-specific pathogenicity in Salmonella.

Animals↗

Predator-induced behavioral and morphological plasticity in the tropical marine Gastropod Strombus gigas.

Florida queen conch stocks once supported a significant fishery, but overfishing prompted the state of Florida to institute a harvest moratorium in 1985. Despite the closure of the fishery, the queen conch population has been slow to recover. One method used in the efforts to restore the Florida conch population has been to release hatchery-reared juvenile conch into the wild; however, suboptimal predator avoidance responses and lighter shell weights relative to their wild counterparts have been implicated in the high mortality rates of released hatchery juveniles. We conducted a series of experiments in which hatchery-reared juvenile conch were exposed to a predator, the spiny lobster (Panulirus argus), to determine whether they could develop behavioral and morphological characteristics that would improve survival. Experiments were conducted in tanks with a calcareous sand substrate to simulate a natural environment. Conditioned conch were exposed to caged lobsters while conch in the control tanks were exposed to empty cages. Conditioned conch moved significantly less and buried themselves more frequently than the naive control conch. Morphometric data indicated that the conditioned conch grew at a significantly slower rate than the naive conch, but the shell weights of the two groups were not significantly different. This implies that the conditioned conch had thicker or denser shells than the control group. As a result, the conditioned conch had significantly higher survival than naive conch in a subsequent predation experiment in which a lobster was allowed to roam free in each tank for 24 hours. In the future, the conditioning protocols documented in this study will be used to increase the survival of hatchery-reared conch in the wild.

Animals↗

Quantification of the spatial aspect of chaotic dynamics in biological and chemical systems.

The need to study spatio-temporal chaos in a spatially extended dynamical system which exhibits not only irregular, initial-value sensitive temporal behavior but also the formation of irregular spatial patterns, has increasingly been recognized in biological science. While the temporal aspect of chaotic dynamics is usually characterized by the dominant Lyapunov exponent, the spatial aspect can be quantified by the correlation length. In this paper, using the diffusion-reaction model of population dynamics and considering the conditions of the system stability with respect to small heterogeneous perturbations, we derive an analytical formula for an 'intrinsic length' which appears to be in a very good agreement with the value of the correlation length of the system. Using this formula and numerical simulations, we analyze the dependence of the correlation length on the system parameters. We show that our findings may lead to a new understanding of some well-known experimental and field data as well as affect the choice of an adequate model of chaotic dynamics in biological and chemical systems.

Animals↗

Individual base model of predator-prey system shows predator dominant dynamics.

Individual base model of predator-prey system is constructed. Both predator and prey species have age structure and cohorts of early reproductive age have competitive advantage. The model has linear functional response in predation behavior and includes the effect of interference among predators and delay of population growth from resource intake, not by functional response but by calculation procedure. Each foraging action is calculated successively and surplus or scarce of acquired resources is interpreted into population size through individual birth and death. This model shows that biomass of prey killed by predator is dependent on demand of predator and that heterogeneity in predator population is essential in persistency and stability of predator-prey system. Heterogeneity of predator makes predator individuals of less competing ability die rapidly. Rapid death of weak individuals causes rapid decrease of total demand of predator and that makes enough room for survived predators. Therefore, the biomass of killed prey is dependent on predator's demand. As young or infant population of predator are the more vulnerable to shortage of prey, and when many of them cannot survive to reproductive age, they can stabilize the system by wasting excessive prey with only temporal numerical increase of predator population.

Animals↗

Responses of Teretrius nigrescens toward the dust and frass of its prey, Prostephanus truncatus.

Teretrius nigrescens is considered to be a specialized predator of Prostephanus truncatus, a serious pest of stored maize and dried cassava roots. Using a bait-bag behavioral bioassay, this investigation found T nigrescens to be strongly arrested by dust and frass produced by P. truncatus on maize, whereas responses to maize flour were weak. Attempts to increase the arrestiveness of flour by altering its physical properties (coarseness or particle size range) were unsuccessful. The arrestive property of dust/frass did not degrade with long-term storage nor did it volatilize or degrade with oven baking up to 150 degrees C. However, extraction with methanol and reapplication was successful in transferring the arrestive property from the dust/frass onto maize flour. The prey dust/frass also induced more oviposition than did maize flour, as judged by production of F1 offspring. The results provide strong evidence for the existence of a high boiling point compound(s) in P. truncatus dust/frass that acts as a contact kairomone for T nigrescens. This, or another compound(s) also seems to act as an oviposition stimulant for female T. nigrescens.

Animals↗

Continuous cycling of grouped vs. solitary strategy frequencies in a predator-prey model.

We present a model of predator and prey grouping strategies using game theory. As predators respond strategically to prey behavior and vice versa, the model is based on a co-evolution approach. Focusing on the "many eyes-many mouths" trade-off, this model considers the benefits and costs of being in a group for hunting predators and foraging prey: predators in a group have more hunting success than solitary predators but they have to share the prey captured; prey in a group face a lower risk of predation but greater competition for resources than lone prey. The analysis of the model shows that the intersections of four curves define distinct areas in the parameter space, corresponding to different strategies used by predators and prey at equilibrium. The model predictions are in accordance with empirical evidence that an open habitat encourages group living, and that low risks of predation favor lone prey. Under some conditions, continuous cycling of the relative frequencies of the different strategies may occur. In this situation, the proportions of grouped vs. solitary predators and prey oscillate over time.

Animals↗