Effects of small-scale habitat fragmentation on predator–prey interactions in a temperate sea grass system

During the last decades, fragmentation has become an important issue in ecological research. Habitat fragmentation operates on spatial scales ranging over several magnitudes from patches to landscapes. We focus on small-scale fragmentation effects relevant to animal foraging decision making that could ultimately generate distribution patterns. In a controlled experimental environment, we tested small-scale fragmentation effects in artificial sea grass on the feeding behaviour of juvenile cod (Gadus morhua). Moreover, we examined the influence of fragmentation on the distribution of one of the juvenile cod’s main prey resources, the grass shrimp (Palaemon elegans), in association with three levels of risk provided by cod (no cod, cod chemical cues and actively foraging cod). Time spent by cod within sea grass was lower in fragmented landscapes, but total shrimp consumption was not affected. Shrimp utilised vegetation to a greater extent in fragmented treatments in combination with active predation. We suggest that shrimp choose between sand and vegetation habitats to minimize risk of predation according to cod habitat-specific foraging capacities, while cod aim to maximize prey-dependent foraging rates, generating a habitat-choice game between predator and prey. Moreover, aggregating behaviour in grass shrimp was only found in treatments with active predation. Hence, we argue that both aggregation and vegetation use are anti-predator defence strategies applied by shrimp. We therefore stress the importance of considering small-scale behavioural mechanisms when evaluating consequences from habitat fragmentation on trophic processes in coastal environments.     

Cannibalism in an age-structured predator–prey system

The effect of cannibalism on an age-structured predator–prey system is studied. Three stable equilibrium states are found. Using a Hopf bifurcation analysis, it is found that the non washout steady state looses its stability as the cannibalism attack rate increases past a bifurcation point S_c. The dependence of the bifurcation point on the other parameters in the model is found. It is shown that the trajectory of the solution spirals in for attack rates S<S_c and exhibits limit cycle behavior for S>S_c.     

Circadian shelter occupancy patterns and predator–prey interactions of juvenile Caribbean spiny lobsters in a reef lagoon

The spiny lobster, Panulirus argus, is predominantly nocturnal, remaining inside shelters during the day and foraging outside at night, presumably to minimize predation risk. Predation risk generally decreases with increasing lobster size. Therefore, this study examined the hypothesis that size would influence this basic circadian pattern. Video cameras continuously recorded the shelter occupancy of juvenile lobsters (n = 72) having a carapace length (CL) of 30–62 mm that were tethered to shelters in a shallow reef lagoon. The lobsters’ shelter occupancy was 100% during the day, but declined linearly from shortly before sunset to a minimum of 50% shortly after midnight and then increased linearly, reaching 100% by 1 h after sunrise. The percent time the lobsters spent in the shelters followed a similar trend, but there was wide variability at night (0–100%) for individual lobsters. Lobsters left their shelters 2–30 times night⁻¹, with a majority of excursions lasting <10 min. These results suggest that juvenile P. argus minimize predation risk by remaining in their shelters as long as possible but offset the energetic cost of this behavior by foraging close to their shelters for several short periods at night. This emergence pattern contrasts with those of early benthic phase lobsters (<15 mm CL), which seldom leave their shelters, and adults (>80 mm CL), which have a dusk/early evening peak in activity and leave the shelter for extended periods of time during the night. Furthermore, a minimum shelter occupancy in the middle of the night appears especially well adapted to avoid exposure to daytime predators. Videotaped observations also included interactions between lobsters and two dominant lobster predators, the triggerfish, Balistes capriscus, and the octopus Octopus cf. vulgaris. Lobsters responded differently to these predators: remaining in the shelter when attacked by a triggerfish and fleeing the shelter when attacked by an octopus. Triggerfish were nearly twice as likely to attack a lobster that was outside of the shelter than inside. Once under attack, however, a lobster had nearly the same chance of surviving if it was inside or outside. Results suggest that the patterns of shelter use and emergence change as lobsters grow, probably reflecting the interplay between perception of predation risk and the need to forage. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Basic features,conjunctive searches,and the confusion effect in predator–prey interactions

The confusion effect describes the observed decrease in the likelihood that a predator will successfully catch any prey when attacking larger groups of moving prey. We introduce readers to the work of cognitive psychologists interested in human visual attention who have been studying their own version of the confusion effect for many years, developing methods and concepts that may be of fundamental utility to behavioral ecologists. In psychology, ‘basic features’ are characteristics unique to a target object in the visual field that no distracter objects share. Images containing targets with basic features are often less likely to induce the confusion effect in human subjects. Target objects with conjunctions of features, on the other hand, have no individual characteristics unique from distracters, but unique characteristics in combination. Such targets more often induce the confusion effect in humans. We propose the ‘basic feature’ (vs. conjunctions of features) as a new organizing concept for studies on the occurrence of the confusion effect in nature, potentially allowing predictions about which types of prey groups are likely to induce the confusion effect in predators.     

Increased nitrogen availability influences predator–prey interactions by altering host-plant quality

Little is known about how plant nutritional and defensive qualities interact to influence predator–prey interactions. To address this need, we provided the neo-tropical milkweed, Asclepias curassavica, with two levels of nitrogen availability and examined how altered host-plant quality influenced the responses of a specialist aphid, Aphis nerii, and a coccinellid predator, Harmonia axyridis. Aphis nerii uses A. curassavica for multiple resources, including nutrition and sequestration of cardenolides for defense against natural enemies. Increased nitrogen availability improved A. curassavica quality by decreasing carbon-to-nitrogen (C:N) ratios and cardenolide concentrations, resulting in A. nerii that also had lower C:N ratios and cardenolide concentrations. Aphis nerii population growth was higher on plants with high nitrogen availability, compared with aphids on plants with low nitrogen availability. In no-choice feeding trials, Harmonia axyridis consumed more high C:N ratio aphids, suggesting a potential compensatory response to reduced aphid nutritional quality. Additionally, H. axyridis were able to consume more low-quality aphids at the expense of increasing exposure to increased cardenolide concentrations, suggesting that interactions between H. axyridis and A. nerii may be strongly influenced by prey nutritional quality. This work highlights the need to consider how variation in plant quality influences herbivore nutritional and defensive quality when examining mechanisms that influence predator–prey interactions.     

Predator–prey models: an application for the plankton dynamics of lake Geneva

In this paper we present a hierarchical Bayesian analysis for a predator–prey model applied to ecology considering the use of Markov Chain Monte Carlo methods. We consider the introduction of a random effect in the model and the presence of a covariate vector. An application to ecology is considered using a data set related to the plankton dynamics of lake Geneva for the year 1990. We also discuss some aspects of discrimination of the proposed models.     

Turning the game around: toxicity in a nudibranch-sponge predator–prey association

Escalation theory proposes enemy-related selection as the most relevant factor of natural selection among individual organisms. When hazardous to predators, prey might be considered enemies that influence predator evolution. Opisthobranch molluscs that prey on chemically defended prey are an interesting study case on this subject. Predation on chemically defended species paved the way for opisthobranchs to enter in an arms race, developing means to detoxify and/or excrete harmful compounds, which led to the sequestration of those compounds and their self-defensive use, an escalation of defenses. Here we aim to understand whether the opisthobranch predator is better protected than its chemically defended prey, using as predator–prey model, a nudibranch (Hypselodoris cantabrica) and the sponge it preys upon (Dysidea fragilis), and from which it obtains deterrent chemical compounds. Specimens of both species were collected on the Portuguese coast, and their crude extracts were analyzed and used in palatability tests. Nudibranchs revealed a higher natural concentration of crude extract, probably due to a progressive accumulation of the compounds. Both predator and prey extracts revealed similar mixtures of deterrent metabolites (furanosesquiterpenes). Palatability tests revealed a more effective deterrence in the nudibranch extracts because significant rejection rates were observed at lower concentrations than those necessary for the sponge extracts to have the same effect. We concluded that the predator is chemically better protected than its prey, which suggests that its acquisition of chemical defenses reveals a defensive escalation.     

Predator–prey fuzzy model

In this work we have used fuzzy rule-based systems to elaborate a predator–prey type of model to study the interaction between aphids (preys) and ladybugs (predators) in citriculture, where the aphids are considered as transmitter agents of the Citrus Sudden Death (CSD). Simulations were performed and a graph was drawn to show the prey population, the potentiality of the predators, and a phase-plane. From this phase-plane, a classic model of the Holling–Tanner type is fitted and its parameters were found. Finally, we have studied the stability of the critical points of the Holling–Tanner model.     

The spider and fly revisited: ploy–counterploy behavior in a unique predator–prey system

Earlier studies have shown that the sarcophagid fly Arachnidomyia lindae is the principal egg-sac predator of the colonial orb-weaving spider Metepeira incrassata, and that risk of egg-sac loss increases with group size. Observations of specialized behaviors for attack (flies) and defense (spiders) suggest that this predator-prey relationship may incorporate elements of ploy and counterploy behavior. Here we explore this relationship in detail and test hypotheses regarding efficacy of attack and defense behaviors. Egg-sac guarding by the spider includes defensive behaviors specific to this fly predator, which were observed during experimental "attacks" with tethered A. lindae, but not with similar presentations of non-predatory Musca domestica. Experimental studies also show that Metepeira incrassata recognizes this predatory fly by airborne cues (i.e., the signature frequency of wing-beats), and can distinguish between this predator and other flies (potential prey) on the basis of wing-beat frequency differences. Removal of female spiders results in a significantly higher probability of unguarded egg-sacs being parasitized, demonstrating the adaptive value of spider defensive behaviors. We also present evidence that A. lindae utilizes a behavioral ploy for circumventing spider guarding behavior (aggressive mimicry - producing vibrations of captured prey in the web), and that Metepeira incrassata, in turn, exhibits a counter-ploy behavior (signal thread cutting) to eliminate this potentially distracting vibratory information. While previous studies have shown that this colonial web-building spider uses a number of general attack-abatement mechanisms against a diversity of predators and parasitoids, results of this study suggest that selection pressures from a highly specialized predator may also result in evolution of predator-specific prey responses.     

Evidence for benthic primary production support of an apex predator–dominated coral reef food web

Five hundred and ninety-nine primary producers and consumers in the Papahānaumokuākea Marine National Monument (PMNM) (22°N–30°N, 160°W–180°W) were sampled for carbon and nitrogen stable isotope composition to elucidate trophic relationships in a relatively unimpacted, apex predator–dominated coral reef ecosystem. A one-isotope (δ¹³C), two-source (phytoplankton and benthic primary production) mixing model provided evidence for an average minimum benthic primary production contribution of 65 % to consumer production. Primary producer δ¹⁵N values ranged from ?1.6 to 8.0 ‰ with an average (2.1 ‰) consistent with a prevalence of N₂ fixation. Consumer group δ¹⁵N means ranged from 6.6 ‰ (herbivore) to 12.1 ‰ (Galeocerdo cuvier), and differences between consumer group δ¹⁵N values suggest an average trophic enrichment factor of 1.8 ‰ Δ¹⁵N. Based on relative δ¹⁵N values, the larger G. cuvier may feed at a trophic position above other apex predators. The results provide baseline data for investigating the trophic ecology of healthy coral reef ecosystems.     

Mechanism of predator–prey detection and behavioral responses in some anuran tadpoles

Predator–prey relationship was studied in three sympatric species of anuran tadpoles. The study design consisted of allowing predaceous Hoplobatrachus tigerinus tadpoles to devour prey tadpoles (Sphaerotheca breviceps and Bufo melanostictus) placed in a plastic tub (five tadpoles of each species, stage ~27) in 30 min. In trials without refugia, more tadpoles of Bufo fell prey compared to Sphaerotheca. In contrast, provision of refugia using hydrilla plant reversed predation risk of the two species. The swimming speed (V _max = 64.55 ± 1.45 cm/s) of Hoplobatrachus tadpoles was much higher compared to the prey species (Bufo: 3.6 ± 0.4 cm/s; Sphaerotheca: 27.6 ± 1.6 cm/s). Poor swimming ability may account for the observed vulnerability of the Bufo tadpoles to predation especially in clear waters; refugia overcame predation to some extent. On the other hand, Sphaerotheca tadpoles that swim faster than the toad tadpoles were less vulnerable in open areas; refugia actually hindered swimming and increased predation. Experiments with association choice tests show that predaceous tadpoles detect prey based on both visual and chemical cues. On the other hand, the prey tadpoles detected predator based exclusively on chemical rather than visual cues. The antipredator defense strategy of the toad tadpoles is manifested in the form of reduced movements, remaining still for longer times and, increased burst speed. The present findings also suggest that in both prey species predator detection has a genetic basis since naive tadpoles with no prior exposure to predators exhibit fright response on first encounter with them.     

Do social spiders cooperate in predator defense and foraging without a web?

Nearly all social spiders spin prey-capture webs, and many of the benefits proposed for sociality in spiders, such as cooperative prey capture and reduced silk costs, appear to depend on a mutually shared web. The social huntsman spider, Delena cancerides (Sparassidae), forms colonies under bark with no capture web, yet these spiders remain in tightly associated, long-lasting groups. To investigate how the absence of the web may or may not constrain social evolution in spiders, we observed D. cancerides colonies in the field and laboratory for possible cooperative defense and foraging benefits. We observed spiders’ responses to three types of potential predators and to prey that were introduced into retreats. We recorded all natural prey capture over 447 h both inside and outside the retreats of field colonies. The colony’s sole adult female was the primary defender of the colony and captured most prey introduced into the retreat. She shared prey with younger juveniles about half the time but never with older subadults. Spiders of all ages individually captured and consumed the vast majority of prey outside the retreat. Young spiders benefited directly from maternal defense and prey sharing in the retreat. However, active cooperation was rare, and older spiders gained no foraging benefit by remaining in their natal colony. D. cancerides does not share many of the benefits of group living described in other web-building social spiders. We discuss other reasons why this species has evolved group living.     

The influence of initial age structure on predator–prey interaction

Initial age structure influences the growth of a prey population and the outcome of the predator–prey interaction. In order to quantify that influence, we employed a simple numerical model using experimental data from the system Tetranychus urticae–Phytoseiulus persimilis. Four major points were drawn from the results: (1) A population created by young females grows much faster than a population created by the same number of females but distributed among the stable age structure. Final number of individuals after a few generations is then much higher than what a plant could support. Consequently, a stable age structure is probably never achieved under these conditions; (2) In the presence of a predator, such a population can persist for a sufficiently long time to overexploit its host plant and to produce enough individuals to allow dispersal; (3) The impact of the predator on the prey population is drastically different according to its own age structure at the beginning of the interaction; and (4) Predators disturb the prey age structure during the course of interactions and thus maintain the prey growth potential at a high level. These points constitute an important adaptation that determine the persistence of the prey and the predator at a metapopulation level. They bring a new insight on the adaptive characters of young female dispersal.     

Biogeochemical transfer and dynamics of iodine in a soil–plant system

Radioactive iodide (¹²⁵I) is used as a tracer to investigate the fate and transport of iodine in soil under various leaching conditions as well as the dynamic transfer in a soil–plant (Chinese cabbage) system. Results show that both soils (the paddy soil and the sandy soil) exhibit strong retention capability, with the paddy soil being slightly stronger. Most iodine is retained by soils, especially in the top 10 cm, and the highest concentration occurs at the top most section of the soil columns. Leaching with 1–2 pore volume water does not change this pattern of vertical distributions. Early breakthrough and long tailing are two features observed in the leaching experiments. Because of the relatively low peak concentration, the early breakthrough is really not an environmental concern of contamination to groundwater. The long tailing implies that the retained iodine is undergoing slow but steady release and the soils can provide a low but stable level of mobile iodine after a short period. The enrichment factors of ¹²⁵I in different plant tissues are ranked as: root > stem > petiole > leaf, and the ¹²⁵I distribution in the young leaves is obviously higher than that in the old ones. The concentrations of ¹²⁵I in soil and Chinese cabbage can be simulated with a dual-chamber model very well. The biogeochemical behaviors of iodine in the soil-cabbage system show that cultivating iodized cabbage is an environmentally friendly and effective technique to eliminate iodine deficiency disorders (IDD). Planting vegetables such as cabbage on the ¹²⁹I-contaminated soil could be a good remediation technique worthy of consideration.     

A predator’s preference for egg-carrying prey: a novel cost of parental care

Using a subsocial spitting spider (Scytodes pallida) as the prey and a spider-eating jumping spider (Portia labiata) as the predator, the cost of parental care is investigated. Our findings suggest that being singled out as preferred prey by P. labiata is, for egg-carrying females of S. pallida, an important cost of parental care. In survival tests, during which P. labiata was given access to egg-carrying and eggless S. pallida females, egg-carrying females were preyed on more often than eggless females. In preference tests, motionless lures instead of living S. pallida were used. The lures were made by mounting dead egg-carrying and dead eggless S. pallida females in lifelike posture in webs. In these tests, P. labiata detected and identified, by vision alone, both kinds of prey (egg-carrying and eggless), and singled out egg-carrying females as preferred prey.Communicated by M.A. Elgar     

Trap barricading and decorating by a well-armored sit-and-wait predator: extra protection or prey attraction?

Animals may build multiple structures to provide benefits to counter the costs of building. Many orb web spiders add multiple structures, e.g., barricading barrier webs and silk decorations, to their webs and these structures have been hypothesized to function to deter predators or attract prey. The heavily armored spiny spiders construct barrier webs around their orb webs and decorate them with conspicuous silk tufts. Why these organisms, already well protected by a thick cuticle and spines, make the extra investment of building barrier webs and adding conspicuous silk decorations is not known. We predicted that these structures function to both attract prey and deter predators. Field experiments were conducted in two consecutive years using orb webs built by the East Asian spiny spider Thelacantha brevispina. We either (1) concealed the decoration, (2) removed the barrier webs, or (3) left the decorations and barrier webs intact. We found year and treatment to interactively influence prey interception rates. In 2010, but not in 2009, we found prey interception with T. brevispina webs to be greater when the decorations were conspicuous than when they were concealed suggesting that the decorations may lure prey. Prey interception was lower when the barrier webs were present without decorations compared to when they were absent without decorations. The prey-attracting function of the decorations thus may counter the reduction in prey interception incurred by adding a barrier web. Predatory wasp interactions were not influenced by any of our treatments, probably because the spiders’ thick cuticle is the primary means of protection from wasps. Bird predation events, while rare, occurred only when decorations were concealed or the barrier webs were removed. It is therefore plausible that the tuft decorations both lure prey and deter birds.     

Fine-scale habitat modeling of a top marine predator: do prey data improve predictive capacity?

Predators and prey assort themselves relative to each other, the availability of resources and refuges, and the temporal and spatial scale of their interaction. Predictive models of predator distributions often rely on these relationships by incorporating data on environmental variability and prey availability to determine predator habitat selection patterns. This approach to predictive modeling holds true in marine systems where observations of predators are logistically difficult, emphasizing the need for accurate models. In this paper, we ask whether including prey distribution data in fine-scale predictive models of bottlenose dolphin (Tursiops truncatus) habitat selection in Florida Bay, Florida, U.S.A., improves predictive capacity. Environmental characteristics are often used as predictor variables in habitat models of top marine predators with the assumption that they act as proxies of prey distribution. We examine the validity of this assumption by comparing the response of dolphin distribution and fish catch rates to the same environmental variables. Next, the predictive capacities of four models, with and without prey distribution data, are tested to determine whether dolphin habitat selection can be predicted without recourse to describing the distribution of their prey. The final analysis determines the accuracy of predictive maps of dolphin distribution produced by modeling areas of high fish catch based on significant environmental characteristics. We use spatial analysis and independent data sets to train and test the models. Our results indicate that, due to high habitat heterogeneity and the spatial variability of prey patches, fine-scale models of dolphin habitat selection in coastal habitats will be more successful if environmental variables are used as predictor variables of predator distributions rather than relying on prey data as explanatory variables. However, predictive modeling of prey distribution as the response variable based on environmental variability did produce high predictive performance of dolphin habitat selection, particularly foraging habitat.     

Fourth-corner correlation is a score test statistic in a log-linear trait–environment model that is useful in permutation testing

Ecologists wish to understand the role of traits of species in determining where each species occurs in the environment. For this, they wish to detect associations between species traits and environmental variables from three data tables, species count data from sites with associated environmental data and species trait data from data bases. These three tables leave a missing part, the fourth-corner. The fourth-corner correlations between quantitative traits and environmental variables, heuristically proposed 20 years ago, fill this corner. Generalized linear (mixed) models have been proposed more recently as a model-based alternative. This paper shows that the squared fourth-corner correlation times the total count is precisely the score test statistic for testing the linear-by-linear interaction in a Poisson log-linear model that also contains species and sites as main effects. For multiple traits and environmental variables, the score test statistic is proportional to the total inertia of a doubly constrained correspondence analysis. When the count data are over-dispersed compared to the Poisson or when there are other deviations from the model such as unobserved traits or environmental variables that interact with the observed ones, the score test statistic does not have the usual chi-square distribution. For these types of deviations, row- and column-based permutation methods (and their sequential combination) are proposed to control the type I error without undue loss of power (unless no deviation is present), as illustrated in a small simulation study. The issues for valid statistical testing are illustrated using the well-known Dutch Dune Meadow data set.     

Incorporating consumer–resource spatial interactions in reserve design

The persistence of species in reserves depends in large part on the persistence of functional ecological interactions. Despite their importance, however, ecological interactions have not yet been explicitly incorporated into conservation prioritization methods. We develop here a general method for incorporating consumer–resource interactions into spatial reserve design. This method protects spatial consumer–resource interactions by protecting areas that maintain the connectivity between the distribution of consumers and resources. We illustrate our method with a conservation planning case study of a mammalian predator, American marten (Martes americana), and its two primary prey species, Red-backed vole (Clethrionomys rutilus) and Deer mouse (Peromyscus maniculatus). The conservation goal was to identify a reserve for marten that comprised 12% of a forest management unit in the boreal forest in Québec, Canada. We compared reserves developed using analysis variants that utilized different levels of information about predator and prey habitat distributions, species-specific connectivity requirements, and interaction connectivity requirements. The inclusion of consumer–resource interactions in reserve-selection resulted in spatially aggregated reserves that maintained local habitat quality for the species. This spatial aggregation was not induced by applying a qualitative penalty for the boundary length of the reserve, but rather was a direct consequence of modelling the spatial needs of the interacting consumer and resources. Our method for maintaining connectivity between consumers and their resources within reserves can be applied even under the most extreme cases of either complete spatial overlap or complete spatial segregation of consumer–resource distributions. The method has been made available via public software.     

Is catchment productivity a useful predictor of taxa richness in lake plankton communities?

The influence of catchment variables on lake organisms is understudied. The terrestrial zone in the vicinity of lakes is, however, probably highly important for biota due to the effects on water chemistry and to various processes operating across ecosystem boundaries. We examined the relative importance of lake and catchment variables, as well as large-scale geographical factors, on the taxa richness of phyto- and zooplankton in 100 small lakes in Finland. In variation partitioning, the variability of phytoplankton richness was most strongly related to the effects of lake variables, the joint effects of lake and catchment variables, and the joint effects of all three groups of variables. Zooplankton richness, in turn, was most strongly related to the effects of lake and catchment variables and the joint effect of lake and catchment variables. The exact results of the variation partitioning depended on the catchment sizes considered in the regression models. Among lake variables, planktonic richness was strongly related to variables indicating productivity. Among catchment variables, the normalized difference vegetation index (NDVI), indicating catchment productivity, showed a relatively strong association with planktonic richness. These results provide evidence that catchment variables such as the NDVI may be efficient predictors of planktonic richness in small lakes. It is possible that individual lakes embedded in a highly productive landscape have higher taxa richness than solitary, potentially productive lakes because of the high influx of dispersing propagules from the regional pool. We also suggest that catchment variables may respond to environmental changes at different scales than the lake variables, and explicit consideration of catchment productivity would therefore be useful when planning research and monitoring programs for freshwater organisms.