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Warning signals and predator-prey coevolution.

Theories of the evolution of warning signals are typically expressed using analytic and computational models, most of which attribute aspects of predator psychology as the key factors facilitating the evolution of warning signals. Sherratt provides a novel and promising perspective with a model that considers the coevolution of predator and prey populations, showing how predators may develop a bias towards attacking cryptic prey in preference to conspicuous prey. Here, we replicate the model as an individual-based simulation and find, in accordance with Sherratt, that predators evolve a bias towards attacking cryptic prey. We then use a Monte Carlo simulation to calculate the relative survivorships of cryptic and conspicuous prey and stress that, as it stands, the model does not predict the evolution or stability of warning signals. We extend the model by giving predators continuous attack strategies and by allowing the evolution of prey conspicuousness: results are robust to the first modification but, in all cases, cryptic prey always enjoy a higher survivorship than conspicuous prey. When conspicuousness is allowed to evolve, prey quickly evolve towards crypsis, even when runaway coevolution is enabled. Sherratt's approach is promising, but other aspects of predator psychology, besides their innate response, remain vital to our understanding of warning signals.

Animals↗

A theoretical investigation of the effect of predators on foraging behaviour and energy reserves.

Data show that when small birds are exposed to a model of a predator, their body mass may either increase or decrease. Although attempts have been made to explain the data using previous models, these models are based on a constant level of predation and hence are not appropriate for making predictions about the response of a bird to the sight of a predator. We have developed a novel model that includes encounters between a bird and potential predators. We show that, depending on the biology of the predator, optimal body mass may either increase or decrease. The model also makes predictions about the foraging behaviour of the bird after it has seen a predator.

Animals↗

Predation risk induces changes in nest-site selection and clutch size in the Siberian jay.

Life-history theory predicts that an individual should reduce its reproductive efforts by laying a smaller clutch size when high risk of nest predation reduces the value of current reproduction. Evidence in favour of this 'nest predation hypothesis', however, is scarce and based largely on correlative analyses. Here, we manipulated perceived risk of nest predation in the Siberian jay Perisoreus infaustus using playback involving a mixture of calls by corvid nest predators in the vicinity of nest sites. In response to being exposed to this acoustic cue simulating increased risk of nest predation, the jays chose a nest site offering more protective covering and reduced clutch size. This is the first experimental demonstration of clutch size adjustment and nest site selection as a result of phenotypic plasticity in an open nesting passerine reflecting a facultative response to the perceived risk of nest predation.

Animals↗

Olfactory assessment of predation risk in the aquatic environment.

The aquatic environment is well suited for the transmission of chemical information. Aquatic animals have evolved highly sensitive receptors for detecting these cues. Here, I review behavioural evidence for the use of chemical cues by aquatic animals for the assessment of predation risk. Chemical cues are released during detection, attack, capture and ingestion of prey. The nature of the cue released depends on the stage of the predation sequence in which cues are released. Predator odours, disturbance pheromones, injury-released chemical cues and dietary cues all convey chemical information to prey Prey use these cues to minimize their probability of being taken on to the next stage of the sequence. The evolution of specialized epidermal alarm substance cells in fishes in the superorder Ostariophysi represent an amplification of this general phenomenon. These cells carry a significant metabolic cost. The cost is offset by the fitness benefit of the chemical attraction of predators. Attempts of piracy by secondary predators interrupt predation events allowing prey an opportunity for escape. In conclusion, chemical cues are widely used by aquatic prey for risk assessment and this has resulted in the evolution of specialized structures among some taxa.

Amphibians↗

Self-organized packs selection in predator-prey ecosystems.

We present a lattice model of a system of predators of five kinds, competing for prey. The predators are grouped in packs and characterized by two parameters-the energy spent on hunting and energy gained by the kill. The success of hunting depends on the actual competition among predators found near a prey. We determine via Monte Carlo simulations the numbers of predators of each kind as a function of time and the distribution of the size of their packs. We show that the ratio of the energy spent by the competing predators determines their fate. The energy gain plays only a secondary role. We show also that the system self-organizes itself into groups of predators living in well defined packs, which size depends on the energy spent. The most preferred size dependence on the energy spent follows a very simple power law. We present also a mean-field-type approach to the problem and we discuss the differences in the results obtained by the two methods, showing in particular, that the simulation approach produces more reliable results.

Animals↗

Phenotypic lability and the evolution of predator-induced plasticity in tadpoles.

The hypothesis that predator-induced defenses in anuran larvae are maintained by divergent selection across multiple predation environments has not been fully supported by empirical results. One reason may be that traits that respond slowly to environmental variation experience a fitness cost not incorporated in the standard adaptive model, due to a time lag between detecting the state of the environment and expressing the phenotypic response. I measured the rate at which behavior and morphology of Rana temporaria tadpoles change when confronted with a switch in the predation environment at two points in development. Hatchling tadpoles that had been exposed during the egg stage to Aeshna dragonfly larvae were not phenotypically different from those exposed as eggs to predator-free conditions, and both responded similarly to post-hatching predator treatments. When 25-day-old tadpoles from treatments with and without dragonflies were subjected to a switch in the environment, their activity budgets reversed completely within 24-36 h, and their body and tail shape began changing significantly within 4 days. The behavioral response was conservative: Tadpoles switched from high-risk to predator-free treatments were slower to adjust their activity. The study confirmed that behavioral traits are relatively labile and exhibit strong plasticity, but it did not reveal such a pattern at the level of individual traits: Morphological traits that developed slowly did not show the least plasticity. Thus, I found that differences in lability of traits were useful for predicting the magnitude of plasticity only for fundamentally different kinds of characters.

Aging↗

From plankton to top predators: bottom-up control of a marine food web across four trophic levels.

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973-2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.

Animals↗

Rapid screening of invertebrate predators for multiple prey DNA targets.

DNA-based techniques are providing valuable new approaches to tracking predator-prey interactions. The gut contents of invertebrate predators can be analysed using species-specific primers to amplify prey DNA to confirm trophic links. The problem is that each predator needs to be analysed with primers for the tens of potential prey available at a field site, even though the mean number of species detected in each gut may be as few as one or two. Conducting all these PCRs (polymerase chain reactions) is a lengthy process, and effectively precludes the analysis of the hundreds of predators that might be required for a meaningful ecological study. We report a rapid, more sensitive and practical approach. Multiplex PCRs, incorporating fluorescent markers, were found to be effective at amplifying degraded DNA from predators' guts and could amplify mitochondrial DNA fragments from 10+ species simultaneously without 'drop outs'. The combined PCR products were then separated by size on polyacrylamide gels on an ABI377 sequencer. New primers to detect the remains of aphids, earthworms, weevils and molluscs in the guts of carabid predators were developed and characterized. The multiplex-sequencer approach was then applied to field-caught beetles, some of which contained DNA from as many as four different prey at once. The main prey detected in the beetles proved to be earthworms and molluscs, although aphids and weevils were also consumed. The potential of this system for use in food-web research is discussed.

Animals↗

Costly carotenoids: a trade-off between predation and infection risk?

Carotenoid reserves in copepods seem costly in terms of predation risk because they make individuals conspicuous. However, carotenoids also seem to play an important role in immune defence as free radical scavengers. To test whether predation risk influences carotenoid levels and whether changes in carotenoid levels are related to changes in immune defence, I examined individual changes in large carotenoid and other lipid droplets upon exposure to predation risk and subsequent exposure to parasites in the copepod Macrocyclops albidus. Copepods reduced carotenoid reserves upon exposure to predators, through which they potentially avoided the costs of being conspicuous under predation risk. Thus, the size of carotenoid reserves is a plastic trait. Such a decrease in carotenoid reserves may also have a negative impact on the copepods' immune system as individuals that decreased their reserves suffered higher parasite prevalence upon exposure to the cestode Schistocephalus solidus. These results suggest that carotenoid reserves may be individually optimized to trade-off each individual's unique costs (predation risk) and benefits (immune defence) of having these reserves.

Animals↗

Delayed effects of larval predation risk and food quality on anuran juvenile performance.

Metamorphosis can disrupt the correlation structure between juvenile and adult traits, thus allowing relatively independent evolution in contrasting environments. We used a multiple experimental approach to investigate how diet quality and larval predation risk affected the rates of growth and development in painted frogs (Discoglossus galganoi), and how these changes influence post-metamorphic performance. A high-energy diet entailed growth advantages only if predation risk did not constrain energy acquisition, whereas diet quality affected primarily the extension of the larval period. Predation risk influenced juvenile shape, most likely via the effects on growth and differentiation rates. Juvenile frogs emerging from predator environments had shorter legs and longer bodies than those from the nonpredator tanks. However, these morphological changes did not translate into differences in relative jumping performance. Neither size-adjusted lipid storage nor fluctuating asymmetry was significantly influenced by food quality or predation risk. Our data suggest that the post-metamorphic costs of predator avoidance during the larval phase are mostly a consequence of small size at metamorphosis.

Animals↗

Size and scaling of predator-prey dynamics.

We propose a scaled version of the Rosenzweig-MacArthur model using both Type I and Type II functional responses that incorporates the size dependence of interaction rates. Our aim is to link the energetic needs of organisms with the dynamics of interacting populations, for which survival is a result of a game-theoretic struggle for existence. We solve the scaled model of predator-prey dynamics and predict population level characteristics such as the scaling of coexistence size ranges and the optimal predator-prey size ratio. For a broad class of such models, the optimal predator-prey size ratio given available prey of a fixed size is constant. We also demonstrate how scaling predictions of prey density differ under resource limitation vs. predator drawdown. Finally, we show how evolution of predator size can destabilize population dynamics, compare scaling of predator-prey cycles to previous work, as well as discuss possible extensions of the model to multispecies communities.

Animals↗

Effects of nutrient availability and Ochromonas sp. predation on size and composition of a simplified aquatic bacterial community.

Predation and competition are two main factors that determine the size and composition of aquatic bacterial populations. Using a simplified bacterial community, composed of three strains characterized by different responses to predation, a short-term laboratory experiment was performed to evaluate adaptations and relative success in communities with experimentally controlled levels of predation and nutrient availability. A strain with a short generation time (Pseudomonas putida), one with high plasticity in cell morphology (Flectobacillus sp. GC5), and one that develops microcolonies (Pseudomonas sp. CM10), were selected. The voracious flagellate Ochromonas sp. was chosen as a predator. To describe adaptations against grazing and starvation, abundance, biomass and relative heterogeneity of bacteria were measured. On the whole, the strains in the predation-free cultures exhibited unicellular growth, and P. putida represented the largest group. The presence of Ochromonas strongly reduced bacterial abundance, but not always the total biomass. The activity of grazers changed the morphological composition of the bacterial communities. Under grazing pressure the relative composition of the community depended on the substrate availability. In the presence of predators, P. putida abundance declined in both high and low nutrient treatments, and Pseudomonas CM10 developed colonies. Flectobacillus was only numerically codominant in the nutrient-rich environments.

Animals↗

Alarm signals of the great gerbil: acoustic variation by predator context, sex, age, individual, and family group.

The great gerbil, Rhombomys opinus, is a highly social rodent that usually lives in family groups consisting of related females, their offspring, and an adult male. The gerbils emit alarm vocalizations in the presence of diverse predators with different hunting tactics. Alarm calls were recorded in response to three predators, a monitor lizard, hunting dog, and human, to determine whether the most common call type, the rhythmic call, is functionally referential with regard to type of predator. Results show variation in the alarm calls of both adults and subadults with the type of predator. Discriminant function analysis classified an average of 70% of calls to predator type. Call variation, however, was not limited to the predator context, because signal structure also differed by sex, age, individual callers, and family groups. These variations illustrate the flexibility of the rhythmic alarm call of the great gerbil and how it might have multiple functions and communicate in multiple contexts. Three alarm calls, variation in the rhythmic call, and vibrational signals generated from foot-drumming provide the gerbils with a varied and multi-channel acoustic repertoire.

Adult↗

Ecological meltdown in predator-free forest fragments.

The manner in which terrestrial ecosystems are regulated is controversial. The "top-down" school holds that predators limit herbivores and thereby prevent them from overexploiting vegetation. "Bottom-up" proponents stress the role of plant chemical defenses in limiting plant depredation by herbivores. A set of predator-free islands created by a hydroelectric impoundment in Venezuela allows a test of these competing world views. Limited area restricts the fauna of small (0.25 to 0.9 hectare) islands to predators of invertebrates (birds, lizards, anurans, and spiders), seed predators (rodents), and herbivores (howler monkeys, iguanas, and leaf-cutter ants). Predators of vertebrates are absent, and densities of rodents, howler monkeys, iguanas, and leaf-cutter ants are 10 to 100 times greater than on the nearby mainland, suggesting that predators normally limit their populations. The densities of seedlings and saplings of canopy trees are severely reduced on herbivore-affected islands, providing evidence of a trophic cascade unleashed in the absence of top-down regulation.

Animals↗

Response in Soil of Cupriavidus necator and Other Copper-Resistant Bacterial Predators of Bacteria to Addition of Water, Soluble Nutrients, Various Bacterial Species, or Bacillus thuringiensis Spores and Crystals.

Soil was incubated with various species of bacteria, Bacillus subtilis, or Bacillus thuringiensis spores and crystals. These were added to serve as potential prey for indigenous, copper-resistant, nonobligate bacterial predators of bacteria in the soil. Alternatively, the soil was incubated with soluble nutrients or water only to cause potential indigenous prey cells to multiply so the predator cells would multiply. All of these incubation procedures caused excessive multiplication of some gram-negative bacteria in soil. Even greater multiplication, however, often occurred for certain copper-resistant bacterial predators of bacteria that made up a part of the gram-negative response. Incubation of the soil with copper per se did not give these responses. In most cases, the copper-resistant bacteria that responded were Cupriavidus necator, bacterial predator L-2, or previously unknown bacteria that resembled them. As was the case for C. necator and L-2, these new bacteria did not use glucose, had white colonies, produced copper-related growth initiation factor (GIF), and attacked B. thuringiensis spores on laboratory media. The results were different, however, when B. thuringiensis spores and crystals per se were added to the soil. The copper-resistant bacterial response in the soil did not, to any extent, include C. necator-like bacteria. Instead, the main copper-resistant bacterial predators that developed had yellow colonies and did not resemble C. necator or L-2 in other ways. They were not seen before, and they did not develop on the addition of B. subtilis spores to soil. Apparently, they could not produce a C. necator-like GIF. Nevertheless, they did respond very quickly to B. thuringiensis spores and crystals in soil, as if a GIF of some sort were involved. These results suggest that, under various conditions of soil incubation, gram-negative bacterial predators of bacteria multiply and that several copper-resistant types among them can be detected, counted, and isolated by plating dilutions of the soil onto media containing excess copper.

Journal Article↗

Role of Pore Size Location in Determining Bacterial Activity during Predation by Protozoa in Soil.

The predation of a luminescence-marked strain of Pseudomonas fluorescens by the soil ciliate Colpoda steinii was studied in soil microcosms. Bacterial cells were introduced in either small (neck diameter, <6 (mu)m) or intermediate-sized (neck diameter, 6 to 30 (mu)m) pores in the soil by inoculation at appropriate matric potentials, and ciliates were introduced into large pores (neck diameter, 30 to 60 (mu)m). Viable cell concentrations of bacteria introduced into intermediate-sized pores decreased at a greater rate than those in small pores, with reductions in bacterial populations being accompanied by an increase in viable cell numbers of the ciliate. The data indicate that the location of bacteria in small pores provides significant protection from predation. In the absence of C. steinii, the level of metabolic activity of the bacterial population, measured by luminometry, decreased at a greater rate than cell number, and the level of luminescence cell(sup-1) consequently decreased. The decrease in levels of luminescence indicates a loss of activity due to starvation. During predation by C. steinii, the level of the activity of cells introduced into small pores fell in a similar manner. The level of cell activity was, however, significantly greater for cells introduced into intermediate-sized pores, despite their greater susceptibility to predation. The data suggest that increased activity arises from a release of nutrients by the predator and the greater accessibility of bacteria to nutrients in larger pores. Nutrient amendment of microcosms resulted in increases in bacterial populations to sustained, higher levels, while levels of luminescence increased transiently. The predation of cells introduced into intermediate-sized pores was greater, and there was also evidence that the level of activity of surviving bacteria was greater for bacteria in intermediate-sized but not small pores.

Journal Article↗

Contrasting bacterial strategies to coexist with a flagellate predator in an experimental microbial assemblage.

We studied predator-induced changes within a slowly growing mixed microbial assemblage that was sustained by algal exudates in a continuous cultivation system. In situ hybridization with fluorescent monolabeled oligonucleotide probes was used for a tentative community analysis. This method also allowed us to quantify the proportions of predators with ingested bacteria of different taxonomic groups. In addition, we determined grazing rates on bacteria with fluorescently labelled prey. Bacteria belonging to the alpha and beta subdivisions of the phylum Proteobacteria ((alpha)- and (beta)-Proteobacteria, respectively) showed very different responses to the addition of a bacterivorous flagellate, Bodo saltans. Within one day, filamentous protist-inedible bacteria developed; these belonged to the (beta)-Proteobacteria and constituted between 8.7 and 34% of bacteria from this subgroup. Total abundance of (beta)-Proteobacteria decreased from 3.05 x 10(sup6) to 0.23 x 10(sup6) cells ml(sup-1), and estimated cell division rates were low. Other morphologically inconspicuous protist-edible bacteria belonging to the (alpha)-Proteobacteria were found to respond to predation by an increase in growth rate. Although these bacteria were heavily grazed upon, as on average >85% of flagellate cells had ingested (alpha)-Proteobacteria, they numerically dominated after the addition of B. saltans (mean, 1.35 x 10(sup6) cells ml(sup-1)). It was thus mainly those fast-dividing strains of (alpha)-Proteobacteria that supported the growth of the flagellate population. We conclude that bacteria in mixed assemblages can adopt at least two distinct strategies as a reaction to intense flagellate predation: to outgrow predation pressure or to develop inedible, inactive filaments. Since these strategies occurred within 24 h after the addition of the flagellate, we hypothesize that chemical stimuli released by the predator may have triggered bacterial responses.

Journal Article↗

Microhabitat preference and vertical use of space by patas monkeys (Erythrocebus patas) in relation to predation risk and habitat structure.

Habitat structure can be important in determining habitat preference of animals because it is often closely linked to factors that affect survival and reproduction, such as food availability and predation risk. Here we examine the ways in which microhabitat structure and predation risk affect the habitat preference of wild patas monkeys (Erythrocebus patas). Patas monkeys in Kenya are typically restricted to Acacia drepanolobium habitat, but within our study group's home range, there are two distinct microhabitats, one with taller trees ('tall microhabitat') and one with apparently perennially shorter trees ('short microhabitat'). Examination of ranging behavior indicates that the patas monkeys preferred the tall microhabitat. In the tall microhabitat, focal animals climbed into trees that were significantly taller than average, indicating that they preferred tall trees. Female patas monkeys spent more time scanning from tall trees than from short trees and detected predators only from taller than average trees, based on alarm call data. Their use of tall trees may have decreased their predation risk by increasing their ability to detect predators. We found no evidence of increased food availability or reduced predator presence in the tall microhabitat that could contribute to the monkeys' preference for the tall microhabitat.

Animals↗