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PubMed1998

Management of fat reserves in tufted titmice Baelophus bicolor in relation to risk of predation.

Fat reserves are an important energy source for animals wintering in temperate zones. Long nights, low ambient temperature and often unpredictable food all increase the probability of death from starvation, and to survive, animals carry energy reserves as fat. Based on the assumption that extra weight makes birds more vulnerable to diurnal predators, it has been hypothesized that predation risk places an upper limit on avian fat reserves. The hypothesis leads to the prediction that birds should decrease their mass in response to increased risk of predation. We tested this prediction with a resident species, the tufted titmouse, Baelophus bicolor. During a 3-day period, we presented a taxidermic mount of a sharp-shinned hawk, Accipiter striatus (predator treatment) or a mourning dove, Zenaida macroura (control treatment) to seven titmice housed individually with food ad libitum. Even though vigilance and delay in time to resume foraging were significantly higher after exposure to the hawk model, the body mass of titmice after exposure to the hawk increased rather than decreased, a result opposite to that predicted. In particular, the birds significantly increased their evening body mass and mean daily mass gain during the treatment period. Our results suggest that risk-dependent foraging appears to regulate avian body fat reserves by limiting foraging time. When food is unlimited, as in our experiment, birds may be able to obtain high energy reserves even while maintaining intense vigilance for predators, and under such conditions, the energetic costs of flying may control upper level energy reserves.Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed1999

Laterality and cooperation: mosquitofish move closer to a predator when the companion is on their left side.

Mirror images simulating social partners that cooperated or defected have been used as an experimental method to test the hypothesis that, while inspecting a predator, pairs of fish play a conditional strategy, Tit for Tat, in an iterated version of the Prisoner's Dilemma game. Using this method, we found that predator inspection was more likely to occur when the mirror image was visible on the left rather than on the right side of mosquitofish, Gambusia holbrooki. The same occurred even when a videorecorded stimulus presentation was used, in which sequences of the predator were mixed with their mirror-image equivalents, thus showing that the asymmetry was not due to behavioural or morphological asymmetries of the predator itself. Moreover, irrespective of whether they were tested with a cooperative (parallel mirror) or a defecting (angled mirror) partner, mosquitofish drew closer to the predator when the mirror was on their left side. These findings suggest that the images seen on the right and left sides by a fish may evoke different types of social behaviour, probably because of differing modes of analysis of perceptual information carried out by the left and right sides of the brain; accurate control and balancing of the side of presentation of visual stimuli during behavioural experiments thus appears to be crucial. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed2000

Decision time and prey gregariousness influence attack probability in naïve and experienced predators.

Aposematic coloration often has an element of conspicuousness. One suggested benefit of conspicuousness is that it enables the prey to be detected at a greater distance, allowing a predator more time to make a correct decision about attacking it and thus reducing possible recognition errors made by predators. I conducted an experiment, with chicks, Gallus gallus domesticus, as predators on live aposematic and nonaposematic prey, to investigate the effects of decision time and signal size on predator sampling behaviour. The chicks were subjected to different degrees of competition to influence how quickly decisions had to be made. Chicks in four treatment groups, either in the presence or absence of a competing chick, were presented with either solitary prey or prey in groups. In the presence of a competitor, chicks attacked the prey more often and more quickly and needed more attacks before they started to avoid the prey. With prey in groups, chicks took longer to attack, attacked less often, learnt to avoid prey more quickly and killed fewer aposematic prey. This experiment provides evidence for the importance of time and signal size for predators' attack decisions. More time to view prey prior to attack could produce a stronger image and thus encourage avoidance learning and produce a stronger neophobic avoidance effect. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed1996

A probabilistic model for deriving soil quality criteria based on secondary poisoning of top predators. II. Calculations for dichlorodiphenyltrichloroethane (DDT) and cadmium.

A simplified food web with three trophic levels is designed: plants and invertebrates at the first, small birds and mammals at the second, and birds and beasts of prey at the third trophic level. Exposure of top predators via separate food chains is analyzed. However, most top predator species are exposed via more than one food chain (food web). Therefore, a species-specific approach is followed too, for which four bird of prey species and two beast of prey species with different food choices are selected: sparrow hawk, kestrel, barn owl, little owl, badger, and weasel. The most critical food chains for secondary poisoning of top predators are soil --> worm/insect --> bird --> bird of prey for dichlorodiphenyltrichloroethane (DDT), and soil --> worm --> bird/mammal --> bird of prey for cadmium (Cd). The risk for the selected top predator species is much lower than the risk based on these critical food chains because the critical food chains constitute a minor part of their food webs. Species feeding on birds (sparrow hawk) and small carnivorous mammals (barn owl) are exposed to DDT and Cd to a much higher extent than species mainly feeding on small herbivorous mammals (kestrel and weasel). It is recommended to include exposure via the pathways soil --> worm/insect --> bird/mammal --> top predator in procedures for derivation of environmental quality objectives for persistent and highly lipophilic compounds.

Animal Feed↗
PubMed2002

Sight of a predator can stimulate a corticosterone response in the great tit (Parus major).

The corticosterone response to the sight of a natural predator was investigated in free-living and captive great tits (Parus major). Free-living great tits responded to the sight of a stuffed, slowly moving Tengmalm's owl, a major predator of great tits, with warning calls and a change in behaviour around a feeder. Great tits returned to the feeder within a few minutes and began to approach the owl, and there was no increase in plasma corticosterone levels in birds sampled 30-50 min after they first saw the owl. Captive great tits in an aviary were exposed for 30 min to a stuffed Tengmalm's owl, to a stuffed brambling, and to a cardboard box. All three stimulus objects were slowly rotated during the exposure period. Great tits exposed to the owl changed their behaviour immediately, and spent most of the time when the owl was visible flying around the aviary and hanging from the roof, with very few visits to a feeder. Great tits exposed to the brambling and to the moving box also changed their behaviour and made fewer visits to the feeder. The great tits responded to the sight of the owl with a marked increase in plasma corticosterone levels, whereas there was no change in corticosterone levels (mean levels < 11 ng/ml) in birds exposed to the brambling or to the moving box. Mean corticosterone levels were high (37.1 +/- 4.9 ng/ml) 0.5 h after exposure to the owl, remained high (38.9 +/- 6.0 ng/ml) 1 h after exposure, and had returned to basal (5.3 plus minus 1.3 ng/ml) 3 h after exposure to the owl. This is the first demonstration for any bird of a complete corticosterone response to a predator. The sight of a predator initiated a corticosterone response in great tits that could not move more than 3 m away, whereas free-living great tits that could choose how far to fly away from the predator either did not initiate a corticosterone response, or had a small corticosterone response in which corticosterone levels were not significantly different from basal 30-50 min later. The results indicate that the initiation of a corticosterone response in birds depends on whether or not a bird perceives that a stimulus is a threat. Furthermore, they illustrate the importance of not making generalised conclusions based on laboratory experiments.

Animals↗
PubMed1995

Evolutionary cycling in predator-prey interactions: population dynamics and the red queen.

This paper describes the coevolution of phenotypes in a community comprising a population of predators and of prey. It is shown that evolutionary cycling is a likely outcome of the process. The dynamical systems on which this description is based are constructed from microscopic stochastic birth and death events, together with a process of random mutation. Births and deaths are caused in part by phenotype-dependent interactions between predator and prey individuals and therefore generate natural selection. Three outcomes of evolution are demonstrated. A community may evolve to a state at which the predator becomes extinct, or to one at which the species coexist with constant phenotypic values, or the species may coexist with cyclic changes in phenotypic values. The last outcome corresponds to a Red Queen dynamic, in which the selection pressures arising from the predator-prey interaction cause the species to evolve without ever reaching an equilibrium phenotypic state. The Red Queen dynamic requires an intermediate harvesting efficiency of the prey by the predator and sufficiently high evolutionary rate constant of the prey, and is robust when the model is made stochastic and phenotypically polymorphic. A cyclic outcome lies outside the contemporary focus on evolutionary equilibria, and argues for an extension to a dynamical framework for describing the asymptotic states of evolution.

Animals↗
PubMed1999

Competition and dispersal in predator-prey waves.

Dispersing predators and prey can exhibit complex spatio-temporal wave-like patterns if the interactions between them cause oscillatory dynamics. We study the effect of these predator-prey density waves on the competition between prey populations and between predator populations with different dispersal strategies. We first describe 1- and 2-dimensional simulations of both discrete and continuous predator-prey models. The results suggest that any population that diffuses faster, disperses farther, or is more likely to disperse will exclude slower diffusing, shorter dispersing, or less likely dispersing populations, everything else being equal. It also appears that it does not matter whether time, space, or state are discrete or continuous, nor what the exact interactions between the predators and prey are. So long as waves exist the competition between populations occurs in a similar fashion. We derive a theory that qualitatively explains the observed behaviour and calculate approximate analytical solutions that describe, to a reasonable extent, these behaviours. Predictions about the cost of dispersal are tested. If strong enough, cost can reverse the populations' relative competitive strengths or lead to coexistence because of the effect of spiral wave cores. The theory is also able to explain previous results of simulations of coexistence in host-parasitoid models (Comins, H. N., and Massell, M. P., 1996, J. Theor. Biol. 183, 19-28).

Animals↗
PubMed2005

Modelling of predator-prey trophic interactions. Part I: two trophic levels.

A class of lumped parameter models to describe the local dynamics in a controlled environment of a two-trophic chain is considered. The class is characterized by a trophic function (functional response of predator to the abundance of prey) depending on the ratio of prey biomass x and a linear function of predator biomass y: f(qx/[(1-rho)k + rhoy]), where q is the efficiency of the predation process, k is a reference biomass, and rho (0 < or = rho < or = 1) specifies the predation model. The trophic function is defined only by some properties determining its shape. A stability analysis of the models has been performed by taking the parameters q and rho as bifurcation parameters: the regions in the (rho,q) plane of existence and stability of nonnegative equilibrium states and limit cycles are determined. This analysis shows that the behaviour of the models is qualitatively similar for 0 < or = rho < 1 (in particular the null state is always a saddle point), while the value rho=1 gives rise to some kind of structural instability of the system (in particular the null state becomes an attractor for sufficiently high predation efficiency).

Animals↗
PubMed2006

Modelling parasitism and predation of mosquitoes by water mites.

Parasitism and predation are two ecological interactions that can occur simultaneously between two species. This is the case of Culicidae (Insecta: Diptera) and water mites (Acari: Hydrachnidia). The larva mites are~parasites of aquatic and semiaquatic insects, and deutonymphs and adults are predators of insect larvae and eggs. Since several families of water mites are associated with mosquitoes there is an interest in the potential use of these mites as biological control agents. The aim of this paper is to use mathematical modelling and analysis to assess the impact of predation and parasitism in the mosquito population. We propose a system of ordinary differential equations to model the interactions among the larval and adult stages of mosquitoes and water mites. The model exhibits three equilibria: the first equilibrium point corresponds to the state where the two species are absent, the second one to the state where only mosquitoes are present (water mites need insects to complete their life cycle), and the third one is the coexistence equilibrium. We analyze conditions for the asymptotic stability of equilibria, supported by analytical and numerical methods. We discuss the different scenarios that appear when we change the parasitism and predation parameters. High rates of parasitism and moderate predation can drive two species to a stable coexistence.

Algorithms↗
PubMed2003

Dryness increases predation risk in efts: support for an amphibian decline hypothesis.

One hypothesis for amphibian declines is that increased dryness attributed to global climate change exposes amphibians to greater biotic threat and, consequently, greater mortality. But, little is known about behavioral responses of terrestrial amphibians to dry conditions alone or in combination with biotic threats. We used field observations and laboratory experiments to test the response of efts (terrestrial juveniles) of the eastern red-spotted newt, Notophthalmus viridescens, to separate and combined desiccation and predation risks. When only at risk of desiccation, efts moved into shade, traveled down slope, decreased activity, and adopted water-conserving postures. Efts also significantly reduced the rate of water loss by huddling and were attracted to chemical cues from conspecific efts but not from conspecific adults. Thus, efts have a variety of behaviors that reduce the risk of dehydration associated with climate change. When faced only with a predation risk, represented by adult and eft newt tissue extracts (alarm chemicals), efts reduced their activity and avoided alarm cues from both sources. When exposed to combined desiccation and predation risks, efts were less active than when exposed to either risk separately and avoided adult tissue extracts, but not eft extracts. These results suggest that under dry conditions, conspecific tissue extracts contain both attractive (huddling) and repulsive (predator-related) chemical components that induce offsetting behavioral responses. This is the first study to demonstrate moisture-dependent responses to conspecific rinses and alarm substances, underscoring the importance of considering environmental moisture and animal hydration in studies examining responses to conspecific odors and/or alarm chemicals. These results support the hypothesis that elevated dehydration risk may compromise anti-predator behavior and exacerbate amphibian population declines.

Analysis of Variance↗
PubMed2003

Prolonged diapause of specialist seed-feeders makes predator satiation unstable in masting of Quercus crispula.

Quercus crispula (= Q. mongolica var. grosseserrata) is the predominant tree species in cool temperate, mixed broadleaf/conifer forests in northern Japan. We compared 11 years of data on acorn production in a population of Q. crispula, with data on seed-insect populations, to try to answer the following questions: (1) Does Q. crispula show a regular pattern of masting? (2) How long do principal seed predators remain in diapause? (3) How do the seed predators affect the pattern of predator satiation? Q. crispula showed a tendency to alternate bearing, with significant synchrony between individual trees. The principal acorn-feeding insects ( Curculio spp. weevils), which infested 25%-70% of matured acorns, generally exhibited a prolonged diapause of 2 years. No significant negative relationship was found between the rate of injury by the weevils and the density of mature acorns, indicating that simple predator satiation fails due to the synchrony of the life-cycle of acorn-feeding insects and the periodical production of acorns. However, the rate of injury by the weevils was negatively correlated with the relative abundance of mature acorns to the number of weevil larvae that had matured 2 years previously. Thus, the proportion of sound acorns notably increased in a rich crop after a disturbance in alternate bearing. Prolonged diapause of specific seed predators is critical in determining the peak year of sound-seed production.

Animals↗
PubMed2003

Damaging UV radiation and invertebrate predation: conflicting selective pressures for zooplankton vertical distribution in the water column of low DOC lakes.

In nature most organisms have to manage conflicting demands of food gathering, predator avoidance, and finding a favorable abiotic environment (oxygen, temperature, etc.) in order to maximize their fitness. In the vertical water column of lakes with high solar ultraviolet radiation (UV) and invertebrate predators, zooplankton face two particularly strong and conflicting selective pressures. During daylight hours invertebrate predators often induce an upward vertical migration of zooplankton prey while potentially damaging UV forces a downward migration. We used 2.2 m long columns suspended vertically in a lake to conduct 2x2 factorial experiments to examine patterns of depth selection behavior by zooplankton in the presence and absence of both the invertebrate predator Chaoborus and UV. We hypothesized that Chaoborus and UV both affect the distribution of zooplankton and a combination of both factors would lead to a narrowing of depth distribution. We found that when Chaoborus were present zooplankton tended to be distributed at shallower depths in the columns, while in the presence of UV they exhibited a deeper distribution. Chaoborus themselves were always found near the bottom of the columns regardless of the UV treatment. Simultaneous exposure to predators and UV resulted in a peak of zooplankton (especially Daphnia catawba) distribution at intermediate depths. In a significant number of cases, depth range was narrowed in response to Chaoborus, UV, or both.

Animals↗
PubMed2004

Movement and spatial organization of small mammals following vertebrate predator exclusion.

Predation directly changes the demographics of prey, generally through a numerical decrease. An indirect effect of predators is alteration of movements and spatial patterns of prey. The relationship between these direct and indirect effects can be tested by excluding predators. Home range size of white-footed mice (Peromyscus leucopus) decreased and home range overlaps increased in the absence of predators. Home range size and overlaps of meadow voles (Microtus pennsylvanicus) did not change despite an increase in vole density. P. leucopus had significantly lower interspecific home range overlaps with M. pennsylvanicus than intraspecific overlaps. The changes in P. leucopus spatial behavior may be an indirect effect of predator exclusion resulting from the increase in M. pennsylvanicus densities.

Analysis of Variance↗
PubMed2004

Influence of recruit condition on food competition and predation risk in a coral reef fish.

Settlement rate is considered to be a major determinant of the population structure of coral reef fishes. In this study, the effects of larval physiological condition on survival, predation risk and competitive ability are assessed for a small damselfish, Pomacentrus moluccensis. New settlers were collected and fed for 5 days to produce high and low condition (measured as lipid) treatment fish. In a field experiment, pairs (one high and one low condition fish) were transplanted to corals. Persistence over 2 weeks was much higher (100% vs. 25%) in high condition fish. In mixed groups in the laboratory, high condition fish were both aggressively dominant and consumed more of a limiting prey source than low condition fish. In addition, low condition fish were shown to be at much higher risk of predation. All of the low condition fish but only 33% of high condition fish in mixed groups were consumed by fish predators, and in a separate experiment, 73% of feeding strikes by predators were directed at low condition fish. Quality of new settlers can have an important influence on subsequent juvenile survival. The mechanisms for this effect are likely to include a combination of effects of condition on food competition and predation risk.

Animals↗
PubMed2004

Rodents balancing a variety of risks: invasive fire ants and indirect and direct indicators of predation risk.

We used foraging trays to compare how oldfield mice, Peromyscus polionotus, altered foraging in response to the presence of fire ants, Solenopsis invicta, and in the presence of direct (predator urine) and indirect (sheltered or exposed microhabitat, moonlight, and precipitation) indicators of predation risk. Foraging reductions elicited by S. invicta were greater than reductions in response to well-documented indicators of risk (i.e., moonlit nights) and the presence of predator urine. The presence of S. invicta always led to reduced foraging, but the overall impact of S. invicta was dependent upon microhabitat and precipitation. When S. invicta was not present, foraging was greater in sheltered microhabitats compared to exposed microhabitats. S. invicta made sheltered microhabitats equivalent to more risky exposed microhabitats, and this effect was especially pronounced on nights without precipitation. The effect of S. invicta suggests that interactions with S. invicta may entail a potentially heavy cost or that presence of S. invicta may represent a more reliable indicator of imminent competition or predation compared to indirect cues of risk and predator urine. The presence of S. invicta led to reduced foraging under situations when foraging activity would otherwise be greatest (i.show $132#e., under vegetative cover), potentially reducing habitat quality for P. polionotus and the distribution of seeds consumed by rodents.

Analysis of Variance↗
PubMed2005

Water turbidity affects predator-prey interactions in a fish-damselfly system.

Community structure may differ dramatically between clear-water and turbid lakes. These differences have been attributed to differences in the cascading effect of fish on prey populations, owing to the reduced efficiency of fish predation in the presence of macrophytes. However, recent theoretical ideas suggest that water turbidity may shape predator-prey interactions, and it is predicted that prey will relax its antipredation behaviour in turbid water (H1). As a result, the nature of predator-prey interactions is expected to shift from both direct and indirect in clear water to dominantly direct in turbid water (H2). We tested these ideas in a fish-damselfly predator-prey system. In a first behavioural experiment, we looked at antipredation behaviour of damselfly larvae isolated from habitats that differ in turbidity, in the presence of fish in clear and turbid water. As predicted in H1, the larvae were more active in turbid than in clear water. In a complementary enclosure experiment, we reared larvae in a clear-water pond and a turbid pond, respectively, and manipulated the origin of the larvae (clear-water, turbid pond), fish presence (absent, present), and vegetation density (sparse, abundant). In both ponds, fish had a direct negative effect on survival of the larvae, which was mitigated in the presence of vegetation. In the fish treatment, the change in average body mass tended to be higher in the turbid pond than in the clear-water pond, suggesting indirect effects of fish were mitigated in the turbid pond. This was supported by a negative effect of fish on the effective growth rate of larvae in the clear pond, but not in the turbid pond. These results are compatible with the idea that predator-prey relationships are mainly governed by direct effects in turbid water, and by direct and indirect effects in clear water.

Analysis of Variance↗
PubMed2005

The role of predation in the decline and extirpation of woodland caribou.

To select appropriate recovery strategies for endangered populations, we must understand the dynamics of small populations and distinguish between the possible causes that drive such populations to low numbers. It has been suggested that the pattern of population decline may be inversely density-dependent with population growth rates decreasing as populations become very small; however, empirical evidence of such accelerated declines at low densities is rare. Here we analyzed the pattern of decline of a threatened population of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Using information on the instantaneous rate of increase relative to caribou density in suitable winter foraging habitat, as well as on pregnancy rates and on causes and temporal distribution of mortalities from a sample of 349 radiocollared animals from 15 subpopulations, we tested 3 hypothesized causes of decline: (a) food regulation caused by loss of suitable winter foraging habitat, (b) predation-sensitive foraging caused by loss of suitable winter foraging habitat and (c) predation with caribou being secondary prey. Population sizes of caribou subpopulations ranged from < 5 to > 500 individuals. Our results showed that the rates of increase of these subpopulations varied from -0.1871 to 0.0496 with smaller subpopulations declining faster than larger subpopulations. Rates of increase were positively related to the density of caribou in suitable winter foraging habitat. Pregnancy rates averaged 92.4% +/-2.24 and did not differ among subpopulations. In addition, we found predation to be the primary cause of mortality in 11 of 13 subpopulations with known causes of mortality and predation predominantly occurred during summer. These results are consistent with predictions that caribou subpopulations are declining as a consequence of increased predation. Recovery of these woodland caribou will thus require a multispecies perspective and an appreciation for the influence of inverse density dependence on population trajectories.

Animals↗
PubMed2005

Size structure and substitutability in an odonate intraguild predation system.

Interactions between different size classes of predator species have the potential to influence survival of prey species in intraguild predation (IGP) systems, but few studies test for these effects. Using a substitutive design in a field setting, I measured the effects of two size classes of IG predators (large and small larvae of the dragonfly Anax junius) on the mortality of IG prey (larvae of the dragonfly Pachydiplax longipennis). I also examined whether combinations of large A. junius and P. longipennis and small A. junius and P. longipennis had substitutable effects on shared prey (larvae of the damselfly Ischnura verticalis). The presence of both size classes of A. junius, when alone and in combination with P. longipennis, significantly increased mortality of I. verticalis. In the presence of P. longipennis, large and small A. junius had similar effects on the mortality of I. verticalis, and effects of size-structured assemblages of A. junius were similar to the effects of each size class alone at the same density. The effects of the two size classes of A. junius on P. longipennis differed, and P. longipennis mortality was lower when exposed to size structured assemblages of A. junius than when exposed to only large A. junius at the same density. Results were similar to those in a laboratory study, although the effect of P. longipennis on I. verticalis was much lower in the field setting. These results demonstrate that interactions between different size classes of IG predators promote the survival of IG prey and highlight the importance of within-species size structure as a characteristic that may promote the coexistence of predators in IGP systems.

Animals↗