Examining intraspecific multiple predator effects across shifting predator sex ratios

Predator-predator interactions, or “multiple predator effects” (MPEs), are pervasive in the structuring of communities and complicate predictive quantifications of ecosystem dynamics. The nature of MPEs is also context-dependent, manifesting differently among species, prey densities and habitat structures. However, there has hitherto been a lack of consideration for the implications of intraspecific demographic variation within populations for the strength of MPEs. The present study extends MPE concepts to examine intraspecific interactions among male and female predators across differences in prey densities using a functional response approach. Focusing on a copepod-mosquito model predator-prey system, interaction strengths of different sex ratio pairs of Lovenula raynerae were quantified towards larval Culex pipiens complex prey, with observations compared to both additive and substitutive model predictions. Copepods exhibited destabilising Type II functional responses in all treatments, with female copepods significantly more voracious than males under multiple predator groups. Lovenula raynerae exhibited significantly negative MPEs overall which resulted in prey risk reductions. However, whilst not statistically clear, the magnitude of antagonistic interactions subtly differed among predator-predator compositions and prey densities, with female-female antagonisms generally prevalent at low prey densities, and male-male negative interactions greater under high prey densities. Mixed-sex copepod group predation was predicted by both additive and substitutive models, and additive models generated significantly higher consumption estimates than substitutive equivalents given that their predictions were based on the absence of antagonistic non-trophic interactions. We propose the importance of internal population sex demographics as a further context-dependency which influences the nature of MPEs, with demographic implications requiring investigation across other taxonomic and trophic groups.     

Seed predator responses

Summary An examination of Enterolobium contortisiliquum predispersal seed predation by beetles suggests that simple tests of the importance of seed density or seed distance from the parent tree must be interpretated with caution.     

Protector turns predator

Therapy-induced autophagy is recognized as a critical determinant of treatment outcome in cancer patients, primarily as a factor underlying drug resistance. However, recent investigations point toward a context-dependent, death-inducing role for autophagy, the mechanism of which remains largely unknown. Our recent study provides evidence that autophagy can directly mediate cell killing in multiple tumor cell types by facilitating degradation of KRAS/K-Ras, a key survival protein. These findings have broad implications for strategies employing autophagy modulation to target tumor cells.     

Influence of predator density on nonindependent effects of multiple predator species

Interactions between multiple predator species are frequent in natural communities and can have important implications for shared prey survival. Predator density may be an important component of these interactions between predator species, as the frequency of interactions between species is largely determined by species density. Here we experimentally examine the importance of predator density for interactions between predator species and subsequent impacts on prey. We show that aggressive interactions between the predatory shore crabs Carcinus maenas and Hemigrapsus sanguineus increased with predator density, yet did not increase as fast as negative interactions between conspecifics. At low density, interactions between conspecific and heterospecific predators had similar inhibitory impacts on predator function, whereas conspecific interference was greater than interference from heterospecifics at high predator density. Thus the impact of conspecific interference at high predator density was sufficient in itself that interactions with a second predator species had no additional impact on per capita predation. Spatial and temporal variability in predator density is a ubiquitous characteristic of natural systems that should be considered in studies of multiple predator species.     

Intraguild predation reduces redundancy of predator species in multiple predator assemblage

1. Interference between predator species frequently decreases predation rates, lowering the risk of predation for shared prey. However, such interference can also occur between conspecific predators. 2. Therefore, to understand the importance of predator biodiversity and the degree that predator species can be considered functionally interchangeable, we determined the degree of additivity and redundancy of predators in multiple- and single-species combinations. 3. We show that interference between two invasive species of predatory crabs, Carcinus maenas and Hemigrapsus sanguineus, reduced the risk of predation for shared amphipod prey, and had redundant per capita effects in most multiple- and single-species predator combinations. 4. However, when predator combinations with the potential for intraguild predation were examined, predator interference increased and predator redundancy decreased. 5. Our study indicates that trophic structure is important in determining how the effects of predator species combine and demonstrates the utility of determining the redundancy, as well as the additivity, of multiple predator species.     

On optimal predator search

Search tactics are explored for animals hunting randomly distributed prey in two-dimensional habitats. The predator chooses between two modes of search: continuous travel and alternating pause-travel. A model based on renewal processes is derived for the predator's net rate of energy intake. The model is used to explore the optimal mode of search, search height, pause duration (“giving-up time”) and move length. Energy expenditure for search is assumed to increase from perchtravel over continuous travel to hover-travel. Prey detectability is assumed to be higher for the pausing than for the travelling predator. The following predictions emerge: The relative merits of the different search modes are mainly determined by the relative magnitudes of energy consumption rates and prey detection efficiencies at pausing and travelling. The energetically cheaper among two search modes reaches its highest relative merit at low prey density and detectability. As either increases, a more expensive search tactic may become superior. If the rate of energy expenditure increases considerably at locomotion, pause-travel tactics may be superior to continuous travel. This requires that the search can be performed from sufficient height, because net energy gain decreases rapidly below the optimum search height. It is greater for pause-travel than for continuous travel, and it increases slightly with decreasing prey density, and markedly with increasing prey detectability. With perch-travel or hover-travel tactics, the optimal giving-up time decreases with increasing prey density and detectability. The optimal move length increases with detectability. Empirical evidence coincides qualitatively with several predictions. Possibilities for further tests of the model are discussed, as are observed behavioural and morphological features on which the search model may shed light.     

Shovelnose sturgeon exhibit predator avoidance behaviour in the presence of a hungry predator

An experiment was designed to test whether age‐0 shovelnose sturgeon (Scaphirhynchus platorynchus) exhibited predator avoidance behaviour in response to a channel catfish (Ictalurus punctatus) predator. It was hypothesized that shovelnose sturgeon would not exhibit any innate predator avoidance behaviour because previous reports have shown a congener of the shovelnose sturgeon, the pallid sturgeon (S. albus), to be an unfavourable prey item for channel catfish. The results, however, indicated that shovelnose sturgeon generally avoided space occupied by the catfish predator and spent a greater proportion of time in the predator avoidance zone within the experimental tank. Bitten fish, in particular, spent a greater period of time in the predator avoidance zone. Of all sturgeon used in this experiment (N = 30), 73% swam within the fork length (350 mm) of the catfish predator. The results seem to indicate that shovelnose sturgeon were initially oblivious to the risk of predation by the catfish predator, but after interaction (e.g. being chased or bitten) appeared to display predator avoidance behaviour. Predator avoidance behaviour in shovelnose sturgeon may thus be suggested as a learned rather than an innate behaviour.     

Global analysis of a predator–prey model with variable predator search rate

We consider a modified Holling-type II predator–prey model, based on the premise that the search rate of predators is dependent on the prey density, rather than constant. A complete analysis of the global behavior of the model is presented, and shows that the model exhibits a dichotomy similar to the classical Holling-type II model: either the coexistence steady state is globally stable; or it is unstable, and then a unique, globally stable limit cycle exists. We discuss the similarities, but also important differences between our model and the Holling-type II model. The main differences are that: 1. The paradox of enrichment which always occurs in the Holling-type II model, does not always occur here, and 2. Even when the paradox of enrichment occurs, predators can adapt by lowering their search rate, and effectively stabilize the system.

     

Neonatal mortality of elk driven by climate, predator phenology and predator community composition

1.?Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. 2.?Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3-5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. 3.?Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). 4.?In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. 5.?Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.     

Prey dispersal and predator persistence

To understand how patchiness influences population dynamics of a tri-trophic interaction, a tractable model is formulated in terms of differential equations. Motivated by the structure of systems such as plants, phytophagous mites and predatory mites, the model takes dispersal into account at the middle trophic level. The effect of dispersal for the middle level in a tri-trophic system could be either stabilising or destabilising since the middle level acts both as prey and as predator. First a simple model with logistic growth for the lowest level is formulated. A model with logistic growth for the lowest level and instantaneous dispersal has a globally stable three-species equilibrium, if this equilibrium exists. Addition of a middle level dispersal phase of non-negligible duration influences equilibrium stability. In the absence of the top trophic level a limit cycle can occur, caused by the delay that exists in the reaction of the middle level to the changes in the lowest level. With low predator efficiency, it is also possible to have an unstable three-species equilibrium. So addition of a middle level dispersal phase of non-negligible duration can work destabilising. Next the persistence of the third trophic level is studied. Even when the three-species equilibrium exists, the third trophic level need not be persistent. A two-species limit cycle can keep its stability when a three-species equilibrium exists; the system is then bistable. It is argued that such a bistability can offer an alternative explanation for pesticide-induced outbreaks of spider mites and failure of predator introduction.     

Nonconsumptive effects in a multiple predator system reduce the foraging efficiency of a keystone predator

Many studies have demonstrated that the nonconsumptive effect (NCE) of predators on prey traits can alter prey demographics in ways that are just as strong as the consumptive effect (CE) of predators. Less well studied, however, is how the CE and NCE of multiple predator species can interact to influence the combined effect of multiple predators on prey mortality. We examined the extent to which the NCE of one predator altered the CE of another predator on a shared prey and evaluated whether we can better predict the combined impact of multiple predators on prey when accounting for this influence. We conducted a set of experiments with larval dragonflies, adult newts (a known keystone predator), and their tadpole prey. We quantified the CE and NCE of each predator, the extent to which NCEs from one predator alters the CE of the second predator, and the combined effect of both predators on prey mortality. We then compared the combined effect of both predators on prey mortality to four predictive models. Dragonflies caused more tadpoles to hide under leaf litter (a NCE), where newts spend less time foraging, which reduced the foraging success (CE) of newts. Newts altered tadpole behavior but not in a way that altered the foraging success of dragonflies. Our study suggests that we can better predict the combined effect of multiple predators on prey when we incorporate the influence of interactions between the CE and NCE of multiple predators into a predictive model. In our case, the threat of predation to prey by one predator reduced the foraging efficiency of a keystone predator. Consequently, the ability of a predator to fill a keystone role could be compromised by the presence of other predators.     

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.     

Breeding synchrony and predator specialization: A test of the predator swamping hypothesis in seabirds

Reproductive synchrony is a widespread phenomenon that is predicted to be adaptive for prey with specialist predators but not for those with generalist ones. I tested this prediction in three polar seabird species characterized by different levels of predator specialization. In the Antarctic petrel, for which the only predator was highly specialized, hatching dates were highly synchronous and chicks that hatched close to the mean hatching date had a higher survival. In black‐legged kittiwakes and Brünnich's guillemots, whose predators were generalists, breeding was less synchronous and there was no fitness advantage in hatching close to the mean. This study emphasizes the potential importance of the relative timing of reproduction for individual fitness and supports the hypothesis that the adaptive value of breeding synchrony depends on the predator functional response.     

Prey kills predator: Counter-attack success of a spider mite against its specific phytoseiid predator

Success of counter-attack by the spider mite,Schizotetranychus celarius (Banks), against its specific phytoseiid predator,Typhlodromus bambusae Ehara, was examined under experimental conditions. The success of counter-attack by prey females (mothers) against a predaceous larva depended upon the former's density per nest. About 30% of the predaceous larvae were killed when they intruded into a nest containing eight females and their offspring. On the other hand, the prey males (fathers) effectively killed the predators, i.e. one male in the nest killed ca. 40% of the predators while two or three males destroyed up to 80%.The presence of prey parents in a nest considerably enhanced the success of the counter-attack. One male and two young females could kill 70% of the predator's larvae, while two males and two females killed 90% of such larvae. This suggests a kind of cooperative brood defence amongstS. celarius parents.Although more robust, protonymphs of the predator also suffered damage by the prey's counter-attack. However, prey male and female could not destroy the predator's eggs and adult females, whilst the latter often killed spider mite adults.From these as well as previous experiments, it is concluded thatS. celarius has evolved some kind of biparental care for its offspring. It is further proposed that the predator—prey interactions observed in this study provide a unique contribution towards understanding predator—prey coevolution.     

Additive effects of predator diversity on pest control caused by few interactions among predator species

1. Studies of the impact of predator diversity on biological pest control have shown idiosyncratic results. This is often assumed to be as a result of differences among systems in the importance of predator–predator interactions such as facilitation and intraguild predation. The frequency of such interactions may be altered by prey availability and structural complexity. A direct assessment of interactions among predators is needed for a better understanding of the mechanisms affecting prey abundance by complex predator communities. 2. In a field cage experiment, the effect of increased predator diversity (single species vs. three‐species assemblage) and the presence of weeds (providing structural complexity) on the biological control of cereal aphids were tested and the mechanisms involved were investigated using molecular gut content analysis. 3. The impact of the three‐predator species assemblages of aphid populations was found to be similar to those of the single‐predator species treatments, and the presence or absence of weeds did not alter the patterns observed. This suggests that both predator facilitation and intraguild predation were absent or weak in this system, or that these interactions had counteracting effects on prey suppression. Molecular gut content analysis of predators provided little evidence for the latter hypothesis: predator facilitation was not detected and intraguild predation occurred at a low frequency. 4. The present study suggests additive effects of predators and, therefore, that predator diversity per se neither strengthens nor weakens the biological control of aphids in this system.     

Intermediate predator impact on consumers weakens with increasing predator diversity in the presence of a top-predator

Adding or removing a top-predator is known to affect lower trophic levels with potentially large, indirect effects on primary production. However, little is known about how predator diversity may affect lower trophic levels, or how adding or removing a top-predator influences the effects of predator diversity. Using aquatic mesocosms containing three and four trophic levels, we tested whether intermediate predator diversity affected predation on consumers and if top-predator presence influenced such effects. We found that the presence of intermediate predators suppressed the consumer population and that this suppression tended to increase with increased intermediate predator diversity when the top-predator was absent. However, with the top-predator present, increased intermediate predator diversity showed the opposite effect on the consumers compared to without a top-predator, i.e. decreased suppression of consumers with increased diversity. Hence, in our study, the loss of intermediate predator species weakened or strengthened predator–prey interactions depending on if the top-predator was present or not, while loss of the top-predator only strengthened the predator–prey interactions. Therefore, the loss of a predator species may render different, but perhaps predictable effects on the functioning of a system depending on from which trophic level it is lost and on the initial number of species in that trophic level.