Testing the threat-sensitive predator avoidance hypothesis: physiological responses and predator pressure in wild rabbits

Predation is a strong selective force with both direct and indirect effects on an animal’s fitness. In order to increase the chances of survival, animals have developed different antipredator strategies. However, these strategies have associated costs, so animals should assess their actual risk of predation and shape their antipredator effort accordingly. Under a stressful situation, such as the presence of predators, animals display a physiological stress response that might be proportional to the risk perceived. We tested this hypothesis in wild European rabbits (Oryctolagus cuniculus), subjected to different predator pressures, in Doñana National Park (Spain). We measured the concentrations of fecal corticosterone metabolites (FCM) in 20 rabbit populations. By means of track censuses we obtained indexes of mammalian predator presence for each rabbit population. Other factors that could modify the physiological stress response, such as breeding status, food availability and rabbit density, were also considered. Model selection based on information theory showed that predator pressure was the main factor triggering the glucocorticoid release and that the physiological stress response was positively correlated with the indexes of the presence of mammalian carnivore predators. Other factors, such as food availability and density of rabbits, were considerably less important. We conclude that rabbits are able to assess their actual risk of predation and show a threat-sensitive physiological response.     

The effects of the functional response on the bifurcation behavior of a mite predator–prey interaction model

 A predator–prey interaction model based on a system of differential equations with temperature-dependent parameters chosen appropriately for a mite interaction on apple trees is analyzed to determine how the type of functional response influences bifurcation and stability behavior. Instances of type I, II, III, and IV functional responses are considered, the last of which incorporates prey interference with predation. It is shown that the model systems with the type I, II, and III functional responses exhibit qualitatively similar bifurcation and stability behavior over the interval of definition of the temperature parameter. Similar behavior is found in the system with the type IV functional response at low levels of prey interference. Higher levels of interference are destabilizing, as illustrated by the prevalence of bistability and by the presence of three attractors for some values of the model parameters. All four systems are capable of modeling population oscillations and outbreaks. Received 12 March 1996; received in revised form 25 October 1996     

Sharks modulate their escape behavior in response to predator size,speed and approach orientation

Escape responses are often critical for surviving predator–prey interactions. Nevertheless, little is known about how predator size, speed and approach orientation impact escape performance, especially in larger prey that are primarily viewed as predators. We used realistic shark models to examine how altering predatory behavior and morphology (size, speed and approach orientation) influences escape behavior and performance in Squalus acanthias, a shark that is preyed upon by apex marine predators. Predator models induced C-start escape responses, and increasing the size and speed of the models triggered a more intense response (increased escape turning rate and acceleration). In addition, increased predator size resulted in greater responsiveness from the sharks. Among the responses, predator approach orientation had the most significant impact on escapes, such that the head-on approach, as compared to the tail-on approach, induced greater reaction distances and increased escape turning rate, speed and acceleration. Thus, the anterior binocular vision in sharks renders them less effective at detecting predators approaching from behind. However, it appears that sharks compensate by performing high-intensity escapes, likely induced by the lateral line system, or by a sudden visual flash of the predator entering their field of view. Our study reveals key aspects of escape behavior in sharks, highlighting the modulation of performance in response to predator approach.     

Prey instar preference and functional responses of Mallada basalis (Walker) (Neuroptera: Chrysopidae) to different life stages of Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae)

Prey instar preference and functional responses of 2- and 3-instar Mallada basalis (Walker) (Neuroptera: Chrysopidae) larvae to 1- to 3-instar Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae) nymphs and adults were assessed in laboratory. Results indicated that both 2- and 3-instar M. basalis larvae preferred young over old P. solenopsis nymphs and adults were the least preferred. The 3-instar M. basalis larvae preyed more adult P. solenopsis than 2-instar larvae. Mallada basalis exhibited type II functional responses to prey densities: An increase in prey density leads to an increase in consumed preys. Regardless of P. solenopsis stages, the number of preys consumed by the 3-instar M. basalis larvae was greater than that by the 2-instar larvae. Attack rates and handling times differed depending on prey and predator stage combinations. The highest attack rate (1.1874) and lowest handling time (0.0040 h) were observed for the 3-instar M. basalis larvae fed on the 1-instar P. solenopsis nymphs. Regardless of P. solenopsis stages, the attack rate of 3-instar M. basalis was greater than 2-instar, whereas the reverse held regarding handling time. The findings collectively indicated that 3-instar M. basalis larvae have greater potential than 2-instar as efficient biological control agent of P. solenopsis. In developing real world biological control programs, however, the 2-instar M. basalis may be released if necessary since the final efficacy of the predator is the summation of the 2- and 3-instar M. basalis.     

Functional and aggregative response of North Sea whiting

The functional response of whiting (Merlangius merlangus L.) to clupeid and gadoid prey was determined from estimates of food intake and prey density at five locations in the North Sea. The intake of most prey types was well described by a type II (decelerating) response, although in some cases a type III (sigmoid) response provided a slightly better fit. Though a saturation level was reached for all types of fish prey, none of the levels corresponded to the maximum digestive capacity of the predator. This was not caused by ingestion of other prey as the amount of other food and fish prey ingested were not negatively correlated. An investigation of the occurrence of fresh fish in the stomachs revealed that fish was ingested almost exclusively during dawn and dusk and the lack of negative correlation between the intake of fish and other prey may thus be a result of the limited time in which fish prey was vulnerable to predation. No aggregative response of the predators was detected towards any of the prey and catches of prey and predators were slightly negatively correlated. There was evidence of an increase in mortality with density at low clupeid densities, but mortality decreased to virtually zero at high densities. Whiting seem therefore unlikely to impose a regulatory effect on their fish prey outside a narrow range of prey densities.     

Assessment of rabbit hemorrhagic disease in controlling the population of red fox: A measure to preserve endangered species in Australia

Predator's management requires a detailed understanding of the ecological circumstances associated with predation. Predation by foxes has been a significant contributor to the Australian native animal reduction. This paper mainly focuses on the dissemination of rabbit hemorrhagic disease in the rabbit population and its subsequences on red fox (Vulpes vulpes) population, by qualitative and quantitative analyses of a designed eco-epidemiological model with simple law of mass action and sigmoid functional response.Existence of solution has been analyzed and shown to be uniformly bounded. The basic reproduction number (R₀) is obtained and the occurrence of a backward bifurcation at R₀ = 1 is shown to be possible using central manifold theory. Global stability of endemic equilibrium is established by geometric approach. Criteria for diffusion-driven ecological instability caused by local random movements of European rabbits and red fox are obtained. Detailed analyses of Turing patterns formation selected by reaction-diffusion system under zero flux boundary conditions are presented. We found that transmission rate, self and cross-diffusion coefficients have appreciable influence on spatial spread of epidemics. Numerical simulation results confirm the analytical finding and generate patterns which indicate that population of red foxes might be controlled if rabbit hemorrhagic disease (RHD) is introduced into the rabbit population and thus ecological balance can be maintained.     

Numerical and functional response of predators to a long-term decline in mammalian prey at a semi-arid Neotropical site

Summary Occurrence and diet of ten carnivorous predators (four falconiforms, four owls, and two foxes), and population levels of their mammalian prey, were monitored over 45 months at a semi-arid site in north-central Chile. Early in this period, small mammals irrupted and then declined markedly to a density 7% of that at peak. All four falconiforms (Buteo polyosoma, Falco sparverius, Geranoaetus melanoleucus, Parabuteo unicinctus) and one owl (Tyto alba) responded numerically to the decline in mammalian prey by virtually abandoning the study site. The three other owls (Athene cunicularia, Bubo virginianus, Glaucidium nanum) and the two foxes (Pseudalopex culpaeus and P. griseus) remained. With few exceptions, throughout the study predators maintained species-specific preferences among small mammal species regardless of the absolute and proportional abundance of these prev. At no time did the two prey species most responsible for the irruption (the rodents Phyllotis darwini and Akodon olivaceus) occur in predators' diets out of proportion to their estimated relative abundance in the field. Predators were clearly unable to prevent the irruption from occurring. Given the absence of a clear functional response to the most irruptive species, predators seemed unlikely to have been responsible for the observed crash. At present, however, predators may be prolonging the crash and delaying the return of small-mammal populations to typical densities.     

An eco-epidemiological system with infected prey and predator subject to the weak Allee effect

In this article, we propose a general prey–predator model with disease in prey and predator subject to the weak Allee effects. We make the following assumptions: (i) infected prey competes for resources but does not contribute to reproduction; and (ii) in comparison to the consumption of the susceptible prey, consumption of infected prey would contribute less or negatively to the growth of predator. Based on these assumptions, we provide basic dynamic properties for the full model and corresponding submodels with and without the Allee effects. By comparing the disease free submodels (susceptible prey–predator model) with and without the Allee effects, we conclude that the Allee effects can create or destroy the interior attractors. This enables us to obtain the complete dynamics of the full model and conclude that the model has only one attractor (only susceptible prey survives or susceptible-infected coexist), or two attractors (bi-stability with only susceptible prey and susceptible prey–predator coexist or susceptible prey-infected prey coexists and susceptible prey–predator coexist). This model does not support the coexistence of susceptible-infected-predator, which is caused by the assumption that infected population contributes less or are harmful to the growth of predator in comparison to the consumption of susceptible prey.     

Experimental removal of piscivorous groupers of the genus Cephalopholis (Serranidae) from coral habitats in the Gulf of Aqaba (Red-Sea)

Synopsis Elimination of adult groupers (Cephalopholis spp.) from various reef formations in the Gulf of Aqaba (Red Sea) resulted in replenishment by other predatory fishes of different taxa and only partly (21.6%) by species of the same genus. Three years later, the initial population composition had not been restored. Recolonization by juvenile groupers presumably settling from the plankton was only 11.3%. Censuses of the typical piscine prey of groupers, before and 36 months after the removal of Cephalopholis, showed virtually no change in abundace and species composition.     

A reappraisal of the evidence for regulation of wolf populations

The dogma that gray wolf (Canis lupus) population densities in naturally occurring systems are limited almost solely by available ungulate biomass is based upon studies that fit straight line linear regressions (Type 1 numerical response) to data collected at 32 sites across North America. We fit Type 1, 2, and 3 response functions to the data using linear and nonlinear regression as appropriate and found that the evidence supported wolf population regulation by density-dependence as much as limitation by prey availability. When we excluded 4 of 32 points from the original data set because those points represented exploited or expanding wolf populations the data suggested that wolf populations are self regulated rather than limited by prey biomass by at least a 3:1 margin. In establishing goals for sustainable wolf population levels, managers of wolf reintroductions and species recovery efforts should account for the possibility that some regulatory mechanism plays an important role in wolf population dynamics. © 2011 The Wildlife Society.     

The myth of constant predator: prey ratios

Apparent constancy in the ratio of predator species to prey species has been offered as evidence that ecological communities are structured by interspecific interactions. If significantly different from random expectation, this effect would be one of the few sound pieces of evidence for community structure. The evidence was re-evaluated by using the data from previous studies to form species pools, and forming simulated communities by drawing species at random from these pools (with replacement). Using a correlation coefficient (number of predator species versus number of prey species), and also the statistic used by the original workers (where different), the observed predator:prey correlation was compared to that for the random communities. In five studies, the observed predator:prey ratio was not significantly different from random expectation. In the only two studies where there was significant departure from the null model, it was with more variation in the ratio than expected on a random basis. It is concluded that there is as yet no evidence for near-constant predator:prey ratios.     

Flavonoids: from cell cycle regulation to biotechnology

Flavonoids have been proposed to play diverse roles in plant growth and development, including defense, symbiosis, pollen development and male fertility, polar auxin transport, and protection against ultraviolet radiation. Recently, a new role in cell cycle regulation has emerged. Genetic alteration of glucuronide metabolism by altered expression of a Pisum sativum UDP-glucuronosyltransferase (PsUGT1) results in an altered cell cycle in pea, alfalfa, and Arabidopsis. In alfalfa, altered expression of PsUGT1 results in accumulation of a flavonoid-like compound that suppresses growth of cultured cells. The results are consistent with the hypothesis that PsUGT1 functions by controlling cellular levels of a factor controlling cell cycle (FCC).     

Tubes and foraging behavior in larval Chironomidae: implications for predator avoidance

Summary In laboratory experiments, I studied differential susceptibility of four co-occurring species of chironomids to a predatory damselfly. The chironomids differed in foraging behavior and could be ranked according to the amount of time they spent outside of their tubes. In choice experiments, the predator consistently selected the prey which spent more time out of the tube, and time out of tube was a significant predictor of the predation rate coefficient. Electivity indices, calculated from field samples and diet analyses of the predator, supported the laboratory results. The data suggest that exposure to predators in a heterogeneous prey community is largely determined by tubedwelling behavior.     

Effects of turbulence on the feeding rate of a pelagic predator: The planktonic hydroid Clytia gracilis

Relatively little is known about the role of turbulence in a predator-prey system where the predator is a passive, pelagic forager. The Campanulariid hydroid Clytia gracilis (Cnidaria, Hydrozoa) is unusual because it occurs as planktonic colonies and is reported to forage passively in the water column on Georges Bank, Massachusetts, USA. In this study, we investigated the role of various turbulence conditions on the feeding rate of C. gracilis colonies in laboratory experiments. We found a positive relationship between turbulence velocities and feeding rates up to a turbulent energy dissipation rate of ca. 1 cm² s^− 3. Beyond this threshold feeding rate decreased slightly, indicating a dome-shaped relationship. Additionally, a negative relationship was found between feeding efficiency and hydroid colony size under lower turbulent velocities, but this trend was not significant under higher turbulence regimes.     

Behavioral responses to prey density by three acarine predator species with different degrees of polyphagy

Behavioral responses by three acarine predators, Phytoseiulus persimilis, Typhlodromus occidentalis, and Amblyseius andersoni (Acari: Phytoseiidae), to different egg and webbing densities of the spider mite Tetranychus urticae (Acari: Tetranychidae) on rose leaflets were studied in the laboratory. Prey patches were delineated by T. urticae webbing and associated kairomones, which elicit turning back responses in predators near the patch edge. Only the presence of webbing affected predator behavior; increased webbing density did not increase patch time. Patch time increased with increased T. urticae egg density in the oligophagous P. persimilis, but was density independent in the polyphagous species T. occidentalis and A. andersoni. Patch time in all three species was more strongly correlated with the number of prey encounters and attacks than with the actual prey number present in the patch. Patch time was determined by (a) the turning back response near the patch edge; this response decayed through time and eventually led to the abandonment of the patch, and (b) encounters with, and attacks upon, prey eggs; these prolonged patch time by both an increment of time spent in handling or rejecting prey and an increment of time spent searching between two successive prey encounters or attacks. Although searching efficiency was independent of prey density in all three species, the predation rate by P. persimilis decreased with prey density because its searching activity (i.e. proportion of total patch time spent in searching) decreased with prey density. Predation rates by T. occidentalis and A. andersoni decreased with prey density because their searching activity and success ratio both decreased with prey density. The data were tested against models of predator foraging responses to prey density. The effects of the degree of polyphagy on predator foraging behavior were also discussed.     

Limits to water transport in Juniperus osteosperma and Pinus edulis: implications for drought tolerance and regulation of transpiration

1. An air-injection method was used to study loss of water transport capacity caused by xylem cavitation in roots and branches of Pinus edulis (Colorado Pinyon) and Juniperus osteosperma (Utah Juniper). These two species characterize the Pinyon–Juniper communities of the high deserts of the western United States. Juniperus osteosperma can grow in drier sites than P. edulis and is considered the more drought tolerant.
2. Juniperus osteosperma was more resistant to xylem cavitation than P. edulis in both branches and roots. Within a species, branches were more resistant to cavitation than roots for P. edulis but no difference was seen between the two organs for J. osteosperma . There was also no difference between juveniles and adults in J. osteosperma ; this comparison was not made for P. edulis .
3. Tracheid diameter was positively correlated with xylem cavitation pressure across roots and stems of both species. This relation suggests a trade-off between xylem conductance and resistance to xylem cavitation in these species.
4. During summer drought, P. edulis maintained higher predawn xylem pressures and showed much greater stomatal restriction of transpiration, consistent with its greater vulnerability to cavitation, than J. osteosperma .
5. These results suggest that the relative drought tolerance of P. edulis and J. osteosperma results in part from difference in their vulnerability to xylem cavitation.     

Piscivore efficiency and refuging prey: the importance of predator search mode

In predator-prey interactions, the efficiency of the predator is dependent on characteristics of both the predator and the prey, as well as the structure of the environment. In a field enclosure experiment, we tested the effects of a prey refuge on predator search mode, predator efficiency and prey behaviour. Replicated enclosures containing young of the year (0+) and 1-year-old (1+) perch were stocked with 3 differentially sized individuals of either of 2 piscivorous species, perch (Perca fluviatilis), pike (Esox lucius) or no piscivorous predators. Each enclosure contained an open predator area with three small vegetation patches, and a vegetated absolute refuge for the prey. We quantified the behaviour of the predators and the prey simultaneously, and at the end of the experiment the growth of the predators and the mortality and habitat use of the prey were estimated. The activity mode of both predator species was stationary. Perch stayed in pairs in the vegetation patches whereas pike remained solitary and occupied the corners of the enclosure. The largest pike individuals stayed closest to the prey refuge whereas the smallest individuals stayed farthest away from the prey refuge, indicating size-dependent interference among pike. Both size classes of prey showed stronger behavioural responses to pike than to perch with respect to refuge use, distance from refuge and distance to the nearest predator. Prey mortality was higher in the presence of pike than in the presence of perch. Predators decreased in body mass in all treatments, and perch showed a relatively stronger decrease in body mass than pike during the experiment. Growth differences of perch and pike, and mortality differences of prey caused by predation, can be explained by predator morphology, predator attack efficiency and social versus interference behaviour of the predators. These considerations suggest that pike are more efficient piscivores around prey refuges such as the littoral zones of lakes, whereas perch have previously been observed to be more efficient in open areas, such as in the pelagic zones of lakes.