The changes in the predatory behavior of the microturbellarian <Emphasis Type="Italic">Stenostomum sphagnetorum</Emphasis> on two species of toxin-secreting ciliates of the genus <Emphasis Type="Italic">Spirostomum</Emphasis>

It is known that the microturbellarian Stenostomum sphagnetorum, a common ciliate predator, is very sensitive against different types of toxins produced by other ciliates for chemical defense, and consequently, it is not able to capture and ingests these ciliates. In particular, when the predator tries to attack one of these toxin-secreting ciliates, it is forced to regurgitate the captured prey in response to the toxin discharged from the ciliates. In this study it is shown that after repeated attacks by S. sphagnetorum against two species of toxic ciliates (Spirostomum ambiguum and Spirostomum teres), the predator acquires a behavior of prey selection that leads to the exclusion of these ciliates among the possible prey and to the distinction between edible and inedible (toxic) ciliates. This learned behavior, which is maintained for days, is lost only after the asexual reproduction of the microturbellarian. In addition, S. sphagnetorum learns to recognize and avoid specimens of S. ambiguum and S. teres artificially deprived of their toxins, strongly suggesting that the toxins are not exclusively associated in the prey recognition of the microturbellarian.     

Tadpole–odonate larvae interactions: influence of body size and diel rhythm

Several studies have shown that prey and predator body size may affect the outcome of predator–prey interactions. However, few studies have taken in account the changes on predator–prey interactions over 24 h. In a tropical freshwater system I evaluated how predator and prey size, and their diel rhythm in activity influenced the interaction between Physalaemus pustulosus tadpoles and dragonfly larvae. Tadpoles of different size classes were exposed to two size classes of the dragonfly larvae Rhionaeschna spec. Feeding trials were conducted during day and night. Tadpole activity showed a diel rhythm and affected size-selective predation of the smallest dragonfly larvae, but not of the larger ones. Predator and prey size had a significant effect on the prey survivorship and prey size had a significant effect on the preference of the predator. The interaction between both factors was significant, indicating that they did not operate independently. I conclude that the predator–prey interactions between odonate larvae and anuran tadpoles were mainly affected by the size of the prey and the predator, and less by the diel activity pattern of the prey.     

Facilitation between herons and smallmouth bass foraging on common prey

Understanding how multiple predators affect one another and their shared prey is an increasingly important goal for ecologists examining predator–prey dynamics and food-web structure. In a field experiment, we examined the outcome of interactions between terrestrial and freshwater predators foraging for the same prey in two temperate North American streams. We used a factorial design to examine the combined foraging effects of herons and smallmouth bass on striped shiners and central stonerollers. We found that there was facilitation between the two predators, resulting in risk-enhancement for the prey species, with particularly pronounced effects on the smallest (<70 mm) size classes. Facilitation is the least well-documented predator–predator interaction and has not been quantitatively demonstrated for freshwater and terrestrial predators. Our results indicate that bass may gain a net benefit from the presence of wading birds such as herons and egrets, and that concerns about the negative effects of birds on fish stocks through competition may be unwarranted.     

Food level and sex shape predator-induced physiological stress: immune defence and antioxidant defence

Despite the potential impact on prey fitness and predator–prey interactions, most studies of predation risk ignore physiological responses and their dependence upon food level and sex. Therefore, we reared male and female larvae of the damselfly Lestes viridis under predator stress (dragonfly larvae) at high and low food levels, and subsequently scored for important variables of insect immune defence (i.e. phenoloxidase) and antioxidant defence [i.e. superoxide dismutase, and catalase (CAT)]. Under predation risk, larvae did not decrease growth rate or immune defence, and only slightly reduced food intake in the high food treatment, probably because of time stress, i.e. little time available to complete the larval development. However, larvae facing predator stress did show an upregulation of antioxidant enzymes. This upregulation was dependent upon food level for CAT and both food level and sex for SOD, consistent with energetic constraints and sex differences in the link between longevity and adult fitness. Our results illustrate that predator stress can influence life history, behavioural and physiological responses differentially and in a context-dependent way. This implies that non-consumptive physiological effects of predators on their prey show independent yet similar complexities in behavioural and life history response variables. In general, our results advocate that mechanistic studies on predator–prey interactions may benefit from including physiological variables.     

Self-optimization, community stability, and fluctuations in two individual-based models of biological coevolution

We compare and contrast the long-time dynamical properties of two individual-based models of biological coevolution. Selection occurs via multispecies, stochastic population dynamics with reproduction probabilities that depend nonlinearly on the population densities of all species resident in the community. New species are introduced through mutation. Both models are amenable to exact linear stability analysis, and we compare the analytic results with large-scale kinetic Monte Carlo simulations, obtaining the population size as a function of an average interspecies interaction strength. Over time, the models self-optimize through mutation and selection to approximately maximize a community potential function, subject only to constraints internal to the particular model. If the interspecies interactions are randomly distributed on an interval including positive values, the system evolves toward self-sustaining, mutualistic communities. In contrast, for the predator–prey case the matrix of interactions is antisymmetric, and a nonzero population size must be sustained by an external resource. Time series of the diversity and population size for both models show approximate 1/f noise and power-law distributions for the lifetimes of communities and species. For the mutualistic model, these two lifetime distributions have the same exponent, while their exponents are different for the predator–prey model. The difference is probably due to greater resilience toward mass extinctions in the food-web like communities produced by the predator–prey model.      

Predator–prey relationships in a Mediterranean vertebrate system: Bonelli’s eagles,rabbits and partridges

How predators impact on prey population dynamics is still an unsolved issue for most wild predator–prey communities. When considering vertebrates, important concerns constrain a comprehensive understanding of the functioning of predator–prey relationships worldwide; e.g. studies simultaneously quantifying ‘functional’ and ‘numerical responses’ (i.e., the ‘total response’) are rare. The functional, the numerical, and the resulting total response (i.e., how the predator per capita intake, the population of predators and the total of prey eaten by the total predators vary with prey densities) are fundamental as they reveal the predator’s ability to regulate prey population dynamics. Here, we used a multi-spatio-temporal scale approach to simultaneously explore the functional and numerical responses of a territorial predator (Bonelli’s eagle Hieraaetus fasciatus) to its two main prey species (the rabbit Oryctolagus cuniculus and the red-legged partridge Alectoris rufa) during the breeding period in a Mediterranean system of south Spain. Bonelli’s eagle responded functionally, but not numerically, to rabbit/partridge density changes. Type II, non-regulatory, functional responses (typical of specialist predators) offered the best fitting models for both prey. In the absence of a numerical response, Bonelli’s eagle role as a regulating factor of rabbit and partridge populations seems to be weak in our study area. Simple (prey density-dependent) functional response models may well describe the short-term variation in a territorial predator’s consumption rate in complex ecosystems.     

Well-informed foraging: damage-released chemical cues of injured prey signal quality and size to predators

Predators use a variety of information sources to locate potential prey, and likewise prey animals use numerous sources of information to detect and avoid becoming the meal of a potential predator. In freshwater environments, chemosensory cues often play a crucial role in such predator/prey interactions. The importance of chemosensory information to teleost fish in marine environments is not well understood. Here, we tested whether coral reef fish predators are attracted to damage-released chemical cues from already wounded prey in order to find patches of prey and minimize their own costs of obtaining food. Furthermore, we tested if these chemical cues would convey information about status of the prey. Using y-maze experiments, we found that predatory dottybacks, Pseudochromis fuscus, were more attracted to skin extracts of damselfish, Pomacentrus amboinensis, prey that were in good condition compared to prey in poor body condition. Moreover, in both the laboratory and field, we found that predators could differentiate between skin extracts from prey based on prey size, showing a greater attraction to extracts made from prey that were the appropriate size to consume. This suggests that predators are not attracted to any general substance released from an injured prey fish instead being capable of detecting and distinguishing relatively small differences in the chemical composition of the skin of their prey. These results have implications for understanding predator foraging strategies and highlights that chemical cues play a complex role in predator–prey interactions in marine fish.     

Influence of ultraviolet-B radiation on growth,prevalence of deformities,and susceptibility to predation in Cascades frog (<Emphasis Type="Italic">Rana cascadae</Emphasis>) larvae

Ambient levels of ultraviolet-B radiation (UVB) have a variety of detrimental effects on aquatic organisms. These include death and effects on growth, development, physiology, and behavior. Amphibians show all of these effects. However, the effects vary with species, life history stage, and ecological context. Little is known about the implications of the detrimental effects of UVB on ecological dynamics. Our study was designed to test how UVB may affect predator–prey interactions, an important ecological dynamic. Specifically, we tested the effect of UVB on the susceptibility of Cascades frog (Rana cascadae) larvae to predation by rough-skinned newts (Taricha granulosa). We also further examined the sublethal effects of UVB on growth and development in Cascades frog larvae. We found no direct effect of UVB exposure on survival. However, UVB-exposed frog larvae displayed decreased growth and increased prevalence of deformities. UVB also caused increased susceptibility to predation, but there was a significant treatment–block interaction. UVB increased susceptibility to predation in two out of five blocks of Cascades frogs. The other three blocks did not show an effect of UVB on susceptibility to predation. Our study suggests that UVB can alter susceptibility to predation in at least one amphibian species. UVB-induced alteration of predator–prey interactions could potentially lead to changes at the population, community, and ecosystem levels. Handling editor: K. Martens     

The chemical ecology of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

We review the chemical ecology of the ladybird beetle Harmonia axyridis from the perspective of its invasiveness and the deleterious effects it exerts in the regions it has colonised. We outline the nature and quantification of its chemical defence, and discuss the protection this provides against natural enemies, particularly intraguild predators. We consider the role of infochemicals in location of prey, intraspecific communication and intraguild interactions. We also discuss the role of prey allelochemicals in relation to H. axyridis extreme dietary generalism. Harmonia axyridis poses a number of practical problems for human health and well-being, including “ladybug taint” wine contamination and problems resulting from large aggregations overwintering in buildings. We consider chemical insights into these issues and, in particular, how attractants and repellents might help manage H. axyridis populations through a push–pull strategy. We conclude by discussing future perspectives for research.     

Identification and characterization of a novel host–toxin interaction in the wheat–<Emphasis Type="Italic">Stagonospora nodorum</Emphasis> pathosystem

Stagonospora nodorum, casual agent of Stagonospora nodorum blotch (SNB) of wheat, produces a number of host-selective toxins (HSTs) known to be important in disease. To date, four HSTs and corresponding host sensitivity genes have been reported, and all four host–toxin interactions are significant factors in the development of disease. Here, we describe the identification and partial characterization of a fifth S. nodorum produced HST designated SnTox4. The toxin, estimated to be 10–30 kDa in size, was found to be proteinaceous in nature. Sensitivity to SnTox4 is governed by a single dominant gene, designated Snn4, which mapped to the short arm of wheat chromosome 1A in a recombinant inbred (RI) population. The compatible Snn4–SnTox4 interaction is light dependent and results in a mottled necrotic reaction, which is different from the severe necrosis that results from other host–toxin interactions in the wheat–S. nodorum pathosystem. QTL analysis in a population of 200 RI lines derived from the Swiss winter wheat varieties Arina and Forno revealed a major QTL for SNB susceptibility that coincided with the Snn4 locus. This QTL, designated QSnb.fcu-1A, explained 41.0% of the variation in disease on leaves of seedlings indicating that a compatible Snn4–SnTox4 interaction plays a major role in the development of SNB in this population. Additional minor QTL detected on the short arms of chromosomes 2A and 3A accounted for 5.4 and 6.0% of the variation, respectively. The effects of the three QTL were largely additive, and together they explained 50% of the total phenotypic variation. These results provide further evidence that host–toxin interactions in the wheat–S. nodorum pathosystem follow an inverse gene-for-gene model.     

A new low cost microbiotest with the freshwater ciliate protozoan Spirostomum ambiguum

This paper reports on preliminary experiments on the controlled culturing of the freshwater ciliated protozoan Spirostomum ambiguum, as a first step in the development of a new low cost microbiotest, to complement the limited battery of standardized cost-effective toxicity tests presently available, with a representative of unicellular biota.Three strains of S. ambiguum originating from France and Poland respectively, have been grown in natural as well as in reconstituted waters, on several types of inert diets and in different temperature and light conditions.The best results in terms of survival and reproduction of the ciliates, leading to a standard recipe for stock culturing, were obtained at 20 °C, in reconstituted Volvic water and on a diet of oat flakes supplemented with dried leave powder as vegetal adjuvant, with weekly renewal of the medium and food. An average generation time of 4 days was obtained in the latter set of stock culturing conditions.     

The response of lions (<Emphasis Type="Italic">Panthera leo</Emphasis>) to changes in prey abundance on an enclosed reserve in South Africa

Large mammalian carnivores place significant pressure on their prey populations and this is exacerbated within the fenced reserves of Africa. However, foraging theory predicts that diet switching by predators may mitigate this pressure. In this study, we use data collected between 2003 and 2007 from an enclosed system in the Eastern Cape Province of South Africa to examine the response of lions Panthera leo to changes in the abundance of two important prey species — kudu Tragelaphus strepsiceros and warthog Phacochoerus africanus. As the relative abundance of warthogs increased, the number of kudu kills decreased significantly, whereas warthog kills became significantly more frequent. A similar pattern was observed for lion prey preference and the switch from kudu to warthog was also reflected in a significant decrease in the mean prey mass. Our results suggest that a diet shift occurs in lions and that the change in diet is primarily in response to an increase in warthog numbers. Prey switching may promote the persistence of predator–prey systems, which is particularly important for fenced systems where natural immigration of prey is not possible. However, continued collection and analysis of long-term observational data from the multipredator, multiprey systems of Africa is required to facilitate a full understanding of predator–prey dynamics.     

Joint evolution of predator body size and prey-size preference

We studied the joint evolution of predator body size and prey-size preference based on dynamic energy budget theory. The predators’ demography and their functional response are based on general eco-physiological principles involving the size of both predator and prey. While our model can account for qualitatively different predator types by adjusting parameter values, we mainly focused on ‘true’ predators that kill their prey. The resulting model explains various empirical observations, such as the triangular distribution of predator–prey size combinations, the island rule, and the difference in predator–prey size ratios between filter feeders and raptorial feeders. The model also reveals key factors for the evolution of predator–prey size ratios. Capture mechanisms turned out to have a large effect on this ratio, while prey-size availability and competition for resources only help explain variation in predator size, not variation in predator–prey size ratio. Predation among predators is identified as an important factor for deviations from the optimal predator–prey size ratio.     

Activity of Sphaerotheca breviceps tadpoles in response to chemical cues of the predaceous tadpoles Hoplobatrachus tigerinus

Tadpoles of Sphaerotheca breviceps raised in the laboratory from the egg stage, and hence lacking prior experience of a predator or its odors, were tested to examine their responses to a predator’s (tadpoles of Hoplobatrachus tigerinus) water-borne chemical cues. The stimulus solution was obtained following 24 h of rearing tadpoles of H. tigerinus (one tadpole per 200 mL water) that were not fed during this period. Upon exposure to the stimulus solution the activity of S. breviceps tadpoles decreased by about 90% within 5 min. Their resting period increased significantly over baseline activity, whereas the swimming period, distance traversed, and swimming spurts declined. However, whenever a test tadpole moved, its swimming velocity was high in response to stimulus solution. The antipredatory responses declined with increase in time of storage of the stimulus solution, indicating decay of the predator’s chemical cues. The findings suggest that (1) antipredator defense strategies of S. breviceps do not require prior experience of predators, (2) the predator’s chemical cues are labile in nature, and (3) the response of prey tadpoles to such cues is similar to reported behavior of anuran tadpoles in response to real predators and alarm cues.     

Self-organised spatial patterns and chaos in a ratio-dependent predator–prey system

Mechanisms and scenarios of pattern formation in predator–prey systems have been a focus of many studies recently as they are thought to mimic the processes of ecological patterning in real-world ecosystems. Considerable work has been done with regards to both Turing and non-Turing patterns where the latter often appears to be chaotic. In particular, spatiotemporal chaos remains a controversial issue as it can have important implications for population dynamics. Most of the results, however, were obtained in terms of ‘traditional’ predator–prey models where the per capita predation rate depends on the prey density only. A relatively new family of ratio-dependent predator–prey models remains less studied and still poorly understood, especially when space is taken into account explicitly, in spite of their apparent ecological relevance. In this paper, we consider spatiotemporal pattern formation in a ratio-dependent predator–prey system. We show that the system can develop patterns both inside and outside of the Turing parameter domain. Contrary to widespread opinion, we show that the interaction between two different type of instability, such as the Turing–Hopf bifurcation, does not necessarily lead to the onset of chaos; on the contrary, the emerging patterns remain stationary and almost regular. Spatiotemporal chaos can only be observed for parameters well inside the Turing–Hopf domain. We then investigate the relative importance of these two instability types on the onset of chaos and show that, in a ratio-dependent predator–prey system, the Hopf bifurcation is indeed essential for the onset of chaos whilst the Turing instability is not.     

Density-dependent effects of prey defences

In this study, we show that the protective advantage of a defence depends on prey density. For our investigations, we used the predator-prey model system Chaoborus-Daphnia pulex. The prey, D. pulex, forms neckteeth as an inducible defence against chaoborid predators. This morphological response effectively reduces predator attack efficiency, i.e. number of successful attacks divided by total number of attacks. We found that neckteeth-defended prey suffered a distinctly lower predation rate (prey uptake per unit time) at low prey densities. The advantage of this defence decreased with increasing prey density. We expect this pattern to be general when a defence reduces predator success rate, i.e. when a defence reduces encounter rate, probability of detection, probability of attack, or efficiency of attack. In addition, we experimentally simulated the effects of defences which increase predator digestion time by using different sizes of Daphnia with equal vulnerabilities. This type of defence had opposite density-dependent effects: here, the relative advantage of defended prey increased with prey density. We expect this pattern to be general for defences which increase predator handling time, i.e. defences which increase attacking time, eating time, or digestion time. Many defences will have effects on both predator success rate and handling time. For these defences, the predator’s functional response should be decreased over the whole range of prey densities. Received: 15 September 1999 / Accepted: 23 December 1999     

Interactions of <Emphasis Type="Italic">Bacillus thuringiensis</Emphasis> Cry1Ac toxin in genetically engineered cotton with predatory heteropterans

A number of cotton varieties have been genetically transformed with genes from Bacillus thuringiensis (Bt) to continuously produce Bt endotoxins, offering whole plant and season-long protection against many lepidopteran larvae. Constant whole-plant toxin expression creates a significant opportunity for non-target herbivores to acquire and bio-accumulate the toxin for higher trophic levels. In the present study we investigated movement of Cry1Ac toxin from the transgenic cotton plant through specific predator-prey pairings, using omnivorous predators with common cotton pests as prey: (1) the beet armyworm, Spodoptera exigua (Lepidoptera: Noctuidae), with the predator Podisus maculiventris (Heteroptera: Pentatomidae); (2) the two-spotted spider mite, Tetranychus urticae (Acarina: Tetranychidae), with the predatory big-eyed bug Geocoris punctipes (Heteroptera: Geocoridae) and (3) with the predatory damsel bug Nabis roseipennis (Heteropera: Nabidae); and (4) the thrips Frankliniella occidentalis (Thysanoptera: Thripidae) with the predatory pirate bug Orius insidiosus (Heteroptera: Anthocoridae). We quantified Cry1Ac toxin in the cotton plants, and in the pests and predators, and the effects of continuous feeding on S. exigua larvae fed either Bt or non-Bt cotton on life history traits of P. maculiventris. All three herbivores were able to convey Cry1Ac toxin to their respective predators. Among the herbivores, T. urticae exhibited 16.8 times more toxin in their bodies than that expressed in Bt-cotton plant, followed by S. exigua (1.05 times), and F. occidentalis immatures and adults (0.63 and 0.73 times, respectively). Of the toxin in the respective herbivorous prey, 4, 40, 17 and 14% of that amount was measured in the predators G. punctipes, P. maculiventris, O. insidiosus, and N. roseipennis, respectively. The predator P. maculiventris exhibited similar life history characteristics (developmental time, survival, longevity, and fecundity) regardless of the prey’s food source. Thus, Cry1Ac toxin is conveyed through non-target herbivores to natural enemies at different levels depending on the herbivore species, but continuous lifetime contact with the toxin by the predator P. maculiventris through its prey had no effect on the predator’s life history. The results found here, supplemented with others already published, suggest that feeding on Cry1Ac contaminated non-target herbivores does not harm predatory heteropterans and, therefore, cultivation of Bt cotton may provide an opportunity for conservation of these predators in cotton ecosystems by reducing insecticide use.     

Prey-Dependent Mortality Rate: A Critical Parameter in Microbial Models

Protozoa are key components of a wide range of ecosystems, but ecological models that incorporate these microbes often suffer from poor parameterisation, specifically of top-level predator loss rates. We (1) suggest that top-level predator mortality is prey-dependent, (2) provide a novel approach to assess this response, and (3) illustrate the ecological relevance of these findings. Ciliates, Paramecium caudatum (prey) and Didinium nasutum (predator), were used to evaluate predator mortality at varying prey levels. To assess mortality, multiple (>100) predators were individually examined (in 2-ml wells), daily (for 3 days), between 0 and 120 preys ml⁻¹. Data were used to determine non-linear mortality and growth responses over a range of prey abundances. The responses, plus literature data were then used to parameterise a predator–prey model, based on the Rosenzweig–MacArthur structure. The model assessed the impact of variable and three levels of constant (high, average and low) mortality rates on P. caudatum–D. nasutum population dynamics. Our method to determine variable mortality rate revealed a strong concave decline in mortality with increasing prey abundance. The model indicated: (1) high- and low-constant mortality rates yielded dynamics that deviate substantially from those obtained from a variable rate; (2) average mortality rate superficially produced dynamics similar to the variable rate, but there were differences in the period of predator–prey cycles, and the lowest abundance of prey and predators (by ~2 orders of magnitude). The differences between incorporating variable and constant mortality rate indicate that including a variable rate could substantially improve microbial-based ecological models.     

Field verification of the use of chemical alarm cues in a coral reef fish

Chemical alarm cues function as early indicators of a predation threat and influence the outcome of predator–prey interactions in the favour of the prey animal. The tropical goby, Asterropteryx semipunctatus, responded with a stereotypical alarm response, including reduced movement and feeding, following exposure to water that contained chemical cues from injured conspecifics under natural field conditions. Gobies did not exhibit an alarm response when challenged with extracts from damaged fish from a different taxonomic family. The behavioural response in the field was similar to that observed in laboratory experiments. This study verifies the use of chemical alarm cues in a marine fish in their natural environment.     

Habitat context influences predator interference interactions and the strength of resource partitioning

Despite increasing evidence that habitat structure can shape predator–prey interactions, few studies have examined the impact of habitat context on interactions among multiple predators and the consequences for combined foraging rates. We investigated the individual and combined effects of stone crabs (Menippe mercenaria) and knobbed whelks (Busycon carica) when foraging on two common bivalves, the hard clam (Mercenaria mercenaria) and the ribbed mussel (Geukensia demissa) in oyster reef and sand flat habitats. Because these species co-occur across these and other estuarine habitats of varying physical complexity, this system is ideal for examining how habitat context influences foraging rates and the generality of predator interactions. Consistent with results from previous studies, consumption rates of each predator in isolation from the other were higher in the sand flat than in the more structurally complex oyster reef habitat. However, consumption by the two predators when combined surprisingly did not differ between the two habitats. This counterintuitive result probably stems from the influence of habitat structure on predator–predator interactions. In the sand-flat habitat, whelks significantly reduced their consumption of their less preferred prey when crabs were present. However, the structurally more complex oyster reef habitat appeared to reduce interference interactions among predators, such that consumption rates when the predators co-occurred did not differ from predation rates when alone. In addition, both habitat context and predator–predator interactions increased resource partitioning by strengthening predator dietary selectivity. Thus, an understanding of how habitat characteristics such as physical complexity influence interactions among predators may be critical to predicting the effects of modifying predator populations on their shared prey.