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PubMed2004

How predation risk affects the temporal dynamics of avian leks: greater sage grouse versus golden eagles.

Leks often attract predators as well as mates, yet most evolutionary models have assumed that sexual selection, not predation, drives lekking behavior. We explored the influence of predation on lek dynamics using a stochastic dynamic game model based on the lek-breeding greater sage grouse (Centrocercus urophasianus) and its principal avian predator, the golden eagle (Aquila chrysaetos). The model predicts time-dependent male lek attendance as a function of factors affecting both mating success (female arrival rate, male numbers, and social status) and predation risk (eagle arrival rate and group size). Dominant males are predicted to arrive sooner and leave later than subordinates, especially if mating skew is high, predation risk is low, or the relationship between lek size and female arrival rate is weak. Both high mean levels of predation risk and small lek size should reduce lek attendance, but the relative tendency of predators to attack large versus small leks has little influence on predicted lekking behavior. Field observations confirmed the predicted effects of female arrival rate, lek size, male dominance, and weather-dependent predator arrival rates on lek departure times. Predicted effects of female arrival rates and male dominance on seasonal lek attendance were also supported. Our model provides an empirically supported adaptive explanation for short-term lek dynamics. It also suggests alternative interpretations for phenomena previously invoked to support the hotshot and skew models of lek formation.

Animals↗
PubMed2006

Habitat selection responses of parents to offspring predation risk: an experimental test.

The ability of nest predation to influence habitat settlement decisions in birds is widely debated, despite its importance in limiting fitness. Here, we experimentally manipulated nest predation risk across a landscape and asked the question, do migratory birds assess and respond to variation in nest predation risk when choosing breeding habitats? We examined habitat preference by quantifying the density and settlement date of eight species of migratory passerines breeding in areas with and without intact nest predator communities. We found consistently more individuals nesting in areas with reduced nest predation than in areas with intact predator assemblages, although predation risk had no influence on settlement or breeding phenology. Additionally, those individuals occupying safer nesting habitats exhibited increased singing activity. These findings support a causal relationship between habitat choice and nest predation risk and suggest the importance of nest predation risk in shaping avian community structure and breeding activity.

Animal Migration↗
PubMed2004

Effect of the red imported fire ant on cotton aphid population density and predation of bollworm and beet armyworm eggs.

The effects of the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), on cotton aphid, Aphis gossypii Glover, populations and its predation of bollworm, Helicoverpa zea (Boddie), and beet armyworm, Spodoptera exigua (Hübner), (both Lepidoptera: Noctuidae) eggs were evaluated in cotton under field conditions during 2001 and 2002 in central and northern Texas. In central Texas, cotton aphid populations were approximately 5.5 times greater and predation of sentinel bollworm eggs 2 times greater in the presence of S. invicta versus in its absence, although aphid populations did not reach economic levels. Most predation of beet armyworm egg masses, measured via direct nocturnal observations, was due to S. invicta (68%) and cotton fleahopper, Pseudatomoscelis seriatus (Reuter) (21%), where S. invicta was present, and by the mite Abrolophus sp. (52%), spiders (13%), and minute pirate bug (Orius sp.) (13%) where S. invicta was absent. Predation of sentinel bollworm eggs and beet armyworm egg masses was approximately 1.5 and 4.1 times greater, respectively, in the presence of S. invicta versus in their absence. In the presence of S. invicta, the relative frequencies of minute pirate bug and cotton fleahopper were higher, and of S. invicta and native ants lower in beat bucket samples compared with their relative frequencies in nocturnal observations of predation upon beet armyworm egg masses. In the absence of S. invicta seven of eight predators sampled were similarly represented in beat bucket samples and nocturnal observations of beet armyworm egg mass predation, whereas minute pirate bug occurred at a higher frequency in beat bucket samples relative to nocturnal observations. These observations suggested that the relative frequencies of minute pirate bug, cotton fleahopper, S. invicta and native ants in beat bucket samples do not closely reflect the frequency with which these predators prey on noctuid eggs. Overall, the results of this study show that although S. invicta may promote aphid populations early in the growing season, it is an important predator of bollworm and beet armyworm eggs later in the season.

Animals↗
PubMed1999

The impact of predation on burrow use by arctic ground squirrels in the boreal forest.

In sedentary animals, the choice of a suitable home site is critical to survival and reproductive fitness. However, habitat suitability may vary with predation risk. We compared habitat use of Arctic ground squirrels (Spermophilus parryii plesius) living in the boreal forest under conditions of fluctuating predation pressure. In our study area, predators show ten-year cycles in numbers that track that of their primary prey, the snowshoe hare (Lepus americanus). In 1993, we compared burrows that continued to be occupied following the period of intense predation during the hare decline of 1990-1992 with those that became vacant, and with random locations. We contrasted these sites to those in a predator exclosure where predation pressure was minimized. Burrows on control sites were located on sloped sites with high visibility. Burrows that remained occupied during the period of intense predation were more likely to be in open areas with fewer fallen trees than burrows that became vacant. We used discriminant functions derived from the control sites and found that 89% of the burrows on the predator exclosure were classified as being similar to the random locations on control sites. We conclude that the distribution of Arctic ground squirrels in the boreal forest is a direct function of predator presence.

Animals↗
PubMed2000

Nest predation increases with parental activity: separating nest site and parental activity effects.

Alexander Skutch hypothesized that increased parental activity can increase the risk of nest predation. We tested this hypothesis using ten open-nesting bird species in Arizona, USA. Parental activity was greater during the nestling than incubation stage because parents visited the nest frequently to feed their young during the nestling stage. However, nest predation did not generally increase with parental activity between nesting stages across the ten study species. Previous investigators have found similar results. We tested whether nest site effects might yield higher predation during incubation because the most obvious sites are depredated most rapidly. We conducted experiments using nest sites from the previous year to remove parental activity. Our results showed that nest sites have highly repeatable effects on nest predation risk; poor nest sites incurred rapid predation and caused predation rates to be greater during the incubation than nestling stage. This pattern also was exhibited in a bird species with similar (i.e. controlled) parental activity between nesting stages. Once nest site effects are taken into account, nest predation shows a strong proximate increase with parental activity during the nestling stage within and across species. Parental activity and nest sites exert antagonistic influences on current estimates of nest predation between nesting stages and both must be considered in order to understand current patterns of nest predation, which is an important source of natural selection.

Animals↗
PubMed2001

Prey scan at random to evade observant predators.

Anti-predator scans by animals occur with very irregular timing, so that the initiation of scans resembles a random, Poisson-like, process. At first sight, this seems both dangerous (predators could exploit the long intervals) and wastefull (scans after very short intervals are relatively uninformative). We explored vigilance timing using a new model that allows both predators and prey to vary their behaviour. Given predators that attack at random with respect to prey behaviour, constant inter-scan intervals minimize predation risk. However, if prey scan regularly to minimize their risk from randomly attacking predators, they become more vulnerable to predators that initiate attacks when the inter-scan intervals begin. If, in order to defeat this tactic, prey choose extremely variable inter-scan intervals, they become more vulnerable to predators who wait for long intervals before launching attacks. Only if predators can monitor the variability of inter-scan intervals and either attack immediately (if variability is too low) or wait for long intervals to attack (if variability is too high) does the empirically observed pattern of Poisson-like scanning become the optimal prey strategy.

Animals↗
PubMed2001

Fat reserves and perceived predation risk in the great tit, Parus major.

The fat reserves of small birds are built up daily as insurance against starvation. They are believed to reflect a trade-off between the risks of starvation and predation such that in situations of high predation risk birds are expected either to reduce their fat reserves in response to mass-dependent predation risk or to increase them in response to foraging interruptions. We assessed the effect on fat reserves of experimentally altering the perceived (but not the actual) risk of predation of wild great tits at a winter feeding site. The perceived predation risk was alternated between 'safe' and 'risky'. Increasing the perceived risk of predation involved 'swooping' a model sparrowhawk over the feeder at four unpredictable times each day using a remote mechanism We produce evidence that the experiment was suceessfull in altering the perceived risk of predation. As predicted from the hypothesis of mass-dependent predation risk, great tits (Parus major) carried significantly reduced fat reserves during the 'risky' treatment. Furthermore, dominant individuals were able to reduce their reserves more than subordinates. As birds returned to feeders within seconds after a predator 'attack', the reduction in fat reserves cannot be attributed to an interruption in feeding.

Adipose Tissue↗
PubMed2004

Predation risk is an ecological constraint for helper dispersal in a cooperatively breeding cichlid.

Environmental conditions are thought to be responsible for the extent and benefits of cooperative breeding in many animal societies, but experimental tests are scarce. We manipulated predator pressure in the cooperatively breeding cichlid Neolamprologus pulcher in Lake Tanganyika, where predators have been suggested to influence helper and breeder survival, helper dispersal and group reproductive success. We varied the type and intensity of predation by releasing medium, large, or no predators inside large underwater cages enclosing two or three group territories. Helper and breeder survival, helper dispersal and group reproductive success decreased from the control, to the medium- and large-predator treatments. These effects were modified by helper body size and the number of adults protecting the group from predators, supporting the 'group augmentation hypothesis'. Predators forced helpers to stay closer to, and spend more time inside, protective shelters. The results demonstrate the importance of predators for group living in this species, and support the 'ecological constraints hypothesis' of cooperative breeding, in the sense that subordinates stay at home rather than leave and breed independently under the risk of predation.

Animals↗
PubMed2006

Multiple predator effects in an intertidal food web.

1. We examined the effects of multiple predators from an intertidal boulder food web to test whether and how three different predator species affected the survival of a small amphipod species. 2. Predators were chosen because they differ in their foraging mode, two feeding at the bottom and in benthic refuges (nemertean and shrimp) and one in the water-column (juveniles of a fish). 3. Mortality of amphipods was not affected by nemerteans, but was high in the presence of shrimp or fish. Highest mortalities were observed in predator-combinations that contained both shrimp and fish. Amphipods responded to shrimp by escaping into the water column, while they avoided fish by remaining in the refuge. We conclude that predator-specific defence causes conflicts for prey when both shrimp and fish are present. 4. Comparing observed effects of multiple predators with expected effects revealed risk enhancement for the shrimp + fish combination. A comparison of different predictive models revealed that the multiplicative model was most appropriate, although additive models may work well under certain conditions. 5. Based on known consumption-ranges of the predators used, we conclude that nemerteans were saturated with prey while fish were far from their saturation point. A predator's functional response curve (prey consumption in relation to prey abundance) determines its impact on prey populations. This knowledge appears essential in order to predict whether prey organisms face risk enhancement, risk reduction or additive effects of multiple predators.

Amphipoda↗
PubMed2006

Frequency-dependent predation and maintenance of prey polymorphism.

In positive frequency-dependent predation, predation risk of an individual prey correlates positively with the frequency of that prey type. In a number of small-scale experiments individual predators have shown frequency-dependent behaviour, often leading to the conclusion that a population of such predators could maintain prey polymorphism. Using simulations, I studied the dynamics of frequency-dependent predation and prey polymorphism. The model suggests that persistence of prey polymorphism decreases with increasing number of predators that show frequency-dependent behaviour, questioning conclusions about polymorphism based on experiments with few predators. In addition, prey population size, prey crypsis, difference in crypsis between prey morphs and the way the behaviour was adjusted affected the persistence of polymorphism. Under some circumstances prey population remained polymorphic for a shorter time under frequency-dependent than under frequency-independent predation. This suggests that although positive frequency-dependent predator behaviour may maintain prey polymorphism, it is not a sufficient condition for persistent prey polymorphism.

Animals↗
PubMed2006

Predation on mutualists can reduce the strength of trophic cascades.

Ecologists have put forth several mechanisms to predict the strength of predator effects on producers (a trophic cascade). We suggest a novel mechanism--in systems in which mutualists of plants are present and important, predators can have indirect negative effects on producers through their consumption of mutualists. The strength of predator effects on producers will depend on their relative consumption of mutualists and antagonists, and on the relative importance of each to producer population dynamics. In a meta-analysis of experiments that examine the effects of predator reduction on the pollination and reproductive success of plants, we found that the indirect negative effects of predators on plants are quite strong. Most predator removal experiments measure the strength of predator effects on producers through the antagonist pathway; we suggest that a more complete understanding of the role of predators will be achieved by simultaneously considering the effects of predators on plant mutualists.

Animals↗
PubMed2006

Escape responses in juvenile Atlantic cod Gadus morhua L.: the effects of turbidity and predator speed.

We examined the effect of turbidity (0.5-14 beam attenuation m(-1)) and predator attack speed (150 and 296 cm s(-1)) on escape responses of juvenile cod Gadus morhua in the laboratory. We triggered escape responses using a predator model and measured escape timing, direction and locomotor performance. We also measured responsiveness and estimated the likelihood of fish escaping the ;predator attack' (putative escape success, PES). Turbidity affected both PES and the type of escape response used by the fish, but these effects depended on predator speed. PES for the fast predator attack declined from 73% in clear water to 21% in highly turbid water, due to decreased responsiveness and poorly timed escapes. Intermediate turbidity enhanced PES and responsiveness to the slow predator attack. Locomotor performance was reduced by turbidity, whereas predator speed had the opposite effect. Our results suggest that both predator attack speed and turbidity have important roles in determining the vulnerability of fish attacked by piscivorous predators.

Animals↗
PubMed1976

The effect of size on the fast-start performance of rainbow trout Salmo cairdneri, and a consideration of piscivorous predator-prey interactions.

The fast-start (acceleration) performance of seven groups of rainbow trout from 9-6 to 38-7 cm total length was measured in response to d.c. electric shock stimuli. Two fast-start kinematic patterns, L- and S-start were observed. In L-starts the body was bent into an L or U shape and a recoil turn normally accompanied acceleration. Free manoeuvre was not possible in L-starts without loss of speed. In S-starts the body was bent into an S-shape and fish accelerated without a recoil turn. The frequency of S-starts increased with size from 0 for the smallest fish to 60-65% for the largest fish. Acceleration turns were common. The radius of smallest turn for both fast-start patterns was proportional to length (L) with an overall radius of 0-17 L. The duration of the primary acceleration stages increased with size from 0-07 s for the group of smallest fish to 0-10 s for the group of largest fish. Acceleration rates were independent of size. The overall mean maximum rate was 3438 cm/s2 and the average value to the end of the primary acceleration movements was 1562 cm/s2. The distance covered and velocity attained after a given time for fish accelerating from rest were independent of size. The results are discussed in the context of interactions between a predator and prey fish following initial approach by the predator. It is concluded that the outcome of an interaction is likely to depend on reaction times of interacting fish responding to manoeuvres initiated by the predator or prey. The prey reaction time results in the performance of the predator exceeding that of the prey at any instant. The predator reaction time and predator error in responses to unpredictable prey manoeuvre are required for prey escape. It is predicted that a predator should strike the prey within 0-1 s if the fish are initially 5-15 cm apart as reported in the literature for predator-prey interactions. These distances would be increased for non-optimal prey escape behaviour and when the prey body was more compressed or depressed than the predator.

Animals↗
PubMed2006

Swordtail fry attend to chemical and visual cues in detecting predators and conspecifics.

Predation pressure and energy requirements present particularly salient opposing selective pressures on young fish. Thus, fry are expected to possess sophisticated means of detecting predators and resources. Here we tested the hypotheses that fry of the swordtail fish Xiphophorus birchmanni use chemical and visual cues in detection of predators and conspecifics. To test these hypotheses we presented young (<7 day-old) fry with combinations of visual and chemical stimuli from adult conspecifics and predators. We found that exposure to predator odors resulted in shoal tightening similar to that observed when fry were presented with visual cues alone. In trials with conspecific stimuli, fry were particularly attracted to adult conspecifics when presented simultaneous visual and chemical stimuli compared to the visual stimulus alone. These results show that fry attend to the odors of adult conspecifics, whose presence in a particular area may signal the location of resources as well as an absence of predators. This is one of the first studies to show that such young fish use chemical and visual cues in predator detection and in interactions with conspecifics. Previous research in X. birchmanni has shown that anthropogenic alteration of the chemical environment disrupts intraspecific chemical communication among adults; we suggest that because fry use the same chemosensory pathways to detect predators and conspecifics, alteration of the chemical environment may critically disrupt predator and resource detection.

Animals↗
PubMed2006

Prey growth and size-dependent predation in juvenile estuarine fishes: experimental and model analyses.

The outcome of predation interactions between growing, size-structured predator and prey cohorts is difficult to predict. We manipulated the food resources available to juvenile spot subject to predation from southern flounder in a 60-day replicated pond experiment to test the hypothesis that spot growing slowly would experience higher predation mortality and stronger selection against small individuals than those growing rapidly. A nearly threefold difference in average growth rate between fast- and slow-growth treatments led to twofold higher predation mortality of slow-growing spot. Relative to no-flounder controls, larger spot were overrepresented at the end of the experiment in both treatments, but the magnitude of flounder size selection was much greater in the slow-growth treatment. The experimental results agreed qualitatively, but not quantitatively, with predictions from a prior size-dependent foraging model. In particular, the model significantly underestimated observed shifts in spot size structure to larger sizes. We hypothesized that competitive release and associated increases in spot growth due to thinning by flounder might reconcile this difference, and extended the model to incorporate this process. We then used the model to estimate the relative contribution of these two confounded predator effects (size-selective predation and thinning) to observed shifts in spot size structure. Model simulations indicated that the combined effects of size-selective predation and thinning could account for nearly all of the observed shift in spot size structure, but that thinning was the more important process. Our results highlight the utility of combining experimental and modeling approaches to unravel the complexities underlying interactions between growing, size-structured predator and prey cohorts.

Animals↗
PubMed2002

Modeling cadmium dynamics in the guts and tissues of small mammals: dose implications for predators.

Predators that consume contaminated small-mammal prey are exposed to pollutants present in both the prey tissues and the prey gut. However, the contribution of the prey gut and its contents to the contaminant intake of the predator is often ignored. This study developed a two-compartment model of the dynamics of Cd in a model prey species, the wood mouse (Apodemus sylvaticus). The results from the Cd dynamics model were used to calibrate a revised predator dose estimation methodology that accounted for contaminant uptake by a predator from both prey tissues and guts. The dose to the predator predicted by the revised methodology was compared to that predicted by conventional dose calculations, which account for contaminants in prey tissues only. The conventional approach underestimated Cd dose to a model predator by a factor of 1.2- to 10-fold, depending on prey age and predator behavior. Thus, accounting for the contaminant burden in the prey gut as well as in the tissues can markedly increase dose estimates to predators, and this is likely to be true for most low-bioavailability pollutants. This study has major implications for ecological risk assessment involving food chain transfer of low-bioavailability xenobiotics to predators.

Animals↗
PubMed2004

Ecological limitations on aquatic mosquito predator colonization in the urban environment.

Urban malaria cases are becoming common in Africa as more people move into cities and industrialization proceeds. While many species of Anopheles mosquitoes vector malaria in rural areas, only a few are found within cities. The success of anthropophilic species in cities, such as members of the An. gambiae complex, may be explained by limitations on colonization by predator species in urban environments. Habitats that are temporal or structurally simple have lower predator survivorship in a variety of ecosystems, but these have not been investigated previously in an urban area. Areas within and around the Kenyan coastal town of Malindi were previously sampled for the presence of standing water using a geographic sampling strategy with probability proportional to size sampling of planned well-drained, unplanned poorly-drained, planned poorly-drained, unplanned well-drained, and peri-urban locations. Standing aquatic habitats in these areas were reassessed. During monthly sampling, presence/absence of mosquitoes and predator taxa were noted, as were ecological habitat variables: structural complexity and presence of water. Lambda statistics were calculated to associate predator guilds, habitat types, location variables, and ecological variables. All predator guilds found in habitats were strongly associated with habitat type, as were the structural complexity and temporal nature of the habitats. Types of habitat were heterogeneously distributed throughout Malindi, with swimming pools as a common habitat type in planned urban areas and tire track pools a common habitat type in peri-urban areas of Malindi. Predator colonization of aquatic habitats in Malindi was strongly influenced by habitat type, and not associated with location characteristics. Ecological variables were affected by the type of habitats, which are co-associated with planning and drainage in Malindi. While habitat types are distributed heterogeneously within Malindi, habitats with low predation pressure are available for mosquito colonization in both urban and peri-urban areas. The temporal, peri-urban tire track pools and the structural simplicity of urban swimming pools may discourage predator colonization, thereby increasing the probability of malaria vectors in these areas of Malindi. Future studies should evaluate habitats for use in malaria surveillance and experimentally test the effects of structural complexity and temporal nature of urban habitats on the densities of mosquito larvae and their aquatic predators.

Animals↗
PubMed1997

Direction of predator approach and the decision to flee to a refuge

How close should an animal allow a potential predator to approach before fleeing to a refuge? Fleeing too soon wastes time and energy that could be spent on other important activities, but fleeing too late is potentially lethal. A model to predict flight initiation distance was developed, based on the assumption that animals would flee at a distance that allows them to reach the refuge ahead of the predator by some margin of safety. This model predicts that (1) flight initiation distance should increase with distance from the refuge (which has been supported by studies on several species) and (2) the rate of increase of flight initiation distance with distance from a refuge should be higher when the refuge is between the predator and prey (prey runs towards the predator) than when the prey is between the predator and the refuge (prey runs away from the predator). Prediction 2 was tested by approaching juvenile woodchucks, Marmota monaxalong an imaginary line between the animal and its burrow entrance and measuring the distance between the observer and the animal at the moment it started its flight. As predicted, the rate of increase in flight initiation distance was higher when the burrow was between the observer and the woodchuck than when the woodchuck was between the observer and the burrow. The slopes were appropriate for predators with pursuit speeds about twice the escape speed of the woodchucks. The difference between the slopes was 1.78 m flight distance/m distance to refuge, close to the value of 2 m flight distance/m distance to refuge predicted by the model. The intercept indicated that woodchucks allowed a margin of safety of about 7.6 m. The model permits quantitative evaluation of the principal elements of flexible escape decisions of animals and provides a measure of how predation risk increases the cost of space use in relation to distance from a refuge.

Journal Article↗