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.     

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.     

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.     

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.     

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     

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.     

Recognising one’s enemies: a functional approach to risk assessment by prey

Little has been done to compare the relative importance of various mechanisms through which prey assess the potential risk from natural enemies. We used predator-naive spider mites (Tetranychus urticae, Tetranychidae) to (1) compare the responses of prey to chemical cues from enemy and non-enemy species and (2) investigate the source of these cues. In the laboratory, we observed the distribution of T. urticae in response to cues from nine mite species, including (1) predators of spider mites, (2) predators/parasites of other animals, and (3) fungivores/pollen-feeders. When given a choice over 24 h, spider mites foraged and oviposited in fewer numbers on leaf discs that were previously exposed to predatory or parasitic mites (including species incapable of attacking spider mites) than on clean leaf discs (unexposed to mites). Interestingly, previous exposure of leaf arenas to fungivores and pollen-feeders had no significant effect on spider mite distribution. We then observed the response of T. urticae to cues from two species of predator that had been reared on a diet of either spider mites or pollen. T. urticae showed stronger avoidance of leaf discs that were previously exposed to spider-mite-fed predators than of discs exposed to pollen-fed predators. Nevertheless, for one predator species (Amblyseius andersoni), T. urticae still preferred to forage and oviposit on clean (unexposed) discs than on discs exposed to pollen-fed predators. Protein-derived metabolic wastes of predatory or parasitic mites may provide a general cue about potential predation risk for T. urticae. However, T. urticae also avoided areas exposed to pollen-fed predators, suggesting there may be other sources of enemy recognition by the spider mites. We discuss the ecological and evolutionary mechanisms that may influence the scope of information through which animals assess predation risk. Received: 11 January 1999 / Received in revised form: 25 October 1999 / Accepted: 20 November 1999     

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.     

Seed choice by rodents: learning or inheritance?

Learning plays a central role in animal life, and it has received special attention in the context of foraging. In this study, we have tested whether learning operates in seed choices by rodents using the Algerian mouse (Mus spretus) and Holm oak (Quercus ilex) acorns as a model. At the laboratory, those rodents captured in the field during the acorn fall period (experienced individuals) rejected weevil-infested acorns, presumably because of their lower energetic value. By contrast, rodents born in captivity and reared without any contact with acorns (naïves) predated weevil-infested acorns at similar rates than sound ones. After exposing naïves to infested and sound acorns during 15 days, they rejected infested acorns as the experienced individuals. In the field, predation on weevil-infested acorns was lower than on sound ones. Predation rates on infested acorns were lowest at the end of the acorn fall season, whereas predation rates of sound acorns increased along the same period. This might be explained by the improved ability to reject infested acorns because of the accumulated experience acquired by the initially naïve rodents along the acorn fall season. We show that learning shapes strongly seed choices by rodents, and it may be advantageous over inherited behaviors in variable unpredictable situations, such as acorn infestation rates that vary strongly between years and trees. We consider that the role of learning has to be taken into account in future studies on seed predation by rodents.     

Nanotechnology in cosmetics: a boon or bane?

Recent advances in the field of nanotechnology enabled the manufacturing of elaborated nanometer-sized particles for various biomedical applications. The cosmetic industry is one of the most enthusiastic early adopters of nanotechnology which routinely uses nano-scale ingredients. However, in the absence of mandatory product labeling, it is difficult to estimate the number of cosmetics, sunscreens and personal care products containing nanoparticles that are now commercially available. The increased capacity of nanoparticles to penetrate skin and gain access to human cells is a double-edged sword as it may be useful for medical purposes, but might also result in far greater uptake of substances that exert adverse health effects. Since nanomaterials represent a large group of structurally, physically, and chemically variable substances, specific toxicological studies are required for each product prior to commercialization. This review aims to delineate the most investigated applications, risk and safety considerations, and regulation of nanotechnology in the field of cosmetics.     

Antarctic shallow-water mega-epibenthos: shaped by circumpolar dispersion or local conditions?

The mega-epibenthos of two different geographic areas, the Antarctic Peninsula and the high Antarctic (eastern Weddell Sea), were investigated using underwater video. The distribution of the marine fauna at shallow depths between 55 and 160 m in these two areas was investigated to determine whether there are any zoogeographic differences at the community level. A total of 237 taxa represented by 85,538 individuals was identified. Multivariate analyses revealed significant faunal differences between northern Marguerite Bay (western Antarctic Peninsula) and the stations from the Weddell Sea, Atka Bay and Four-Seasons Bank. Echinoderms, especially ophiuroids, dominated Marguerite Bay, bryozoans and ascidians were abundant at Atka Bay, and hydroids and gorgonians were well represented at Four-Seasons Bank. These clear differences can mainly be explained by the influence of local environmental conditions that are probably the primary feature responsible in shaping the Antarctic shallow-water epifauna and not an intensive exchange with larger depths or a limited dispersion due to scarce and isolated shallow areas. In addition, modes of reproduction and characteristics of the early life history (e.g. brooding, viviparity or budding) of key taxa may also shape patterns of species distribution in shallow benthic Antarctic communities.Communicated by O. Kinne, Oldendorf/Luhe     

Localized defecation by pike: a response to labelling by cyprinid alarm pheromone?

Fathead minnows (Pimephales promelas) that have never encountered a predatory pike (Esox lucius), are able to detect conspecific alarm pheromone in a pike's diet if the pike has recently consumed minnows. It remains unclear how this minnow alarm pheromone is secreted by pike and if a pike is able to avoid being labelled as a potential predator by localizing these cues away from its foraging range. The first experiment determined that minnow alarm pheromone is present in pike feces when pike are fed minnows. Individual fathead minnows exhibited a fright response to a stimulus of pike feces if the pike had been fed minnows, but not if the pike had been fed swordtails, which lack alarm pheromone. Individual minnows also exhibited a fright reaction to alarm pheromone in the water (which contained no feces) housing pike which had been fed minnows, suggesting that alarm pheromone is also released in urine, mucous secretions and/or via respiration. The second experiment determined that test pike spent a significantly greater proportion of time in the home area of the test tanks (i.e. where they were fed) but the majority of feces were deposited in the opposite end of the test tank. By localizing their defecation away from the home or foraging area, pike may be able to counter the effects of being labelled as a predator by the alarm pheromone of the prey species.     

A predator’s body coloration enhances its foraging profitability by day and night

Few predators forage by both day and night. It remains unknown, however, how the costs and benefits of foraging or signaling are partitioned in animals that forage at all times. The orb-web spider Cyrtophora moluccensis is brightly colored and forages by day and night. We determined the benefits reaped when it forages by both day and night by estimating the biomass of prey caught in their webs. Additionally, we quantified whether the spider’s presence influences the number of prey caught by day and night and whether its colorful body is visible to diurnal and/or nocturnal insects using diurnal and nocturnal insect vision models. We found that approximately five times the biomass of prey was caught in C. moluccensis’ webs by night than by day. Hemipterans, hymenopterans, and dipterans were predominantly caught by day, while lepidopterans (moths) were predominately caught by night. Accordingly, we concluded that foraging by night is more profitable than foraging by day. We predicted that other benefits, for example, energetic advantages or enhanced fecundity, may promote its daytime activity. Foraging success was greater by day and night when the spider was present in the web than when the spider was absent. We also found that parts of the spider’s body were conspicuous to diurnal and nocturnal insects, possibly through different visual channels. The colorful body of C. moluccensis, accordingly, appears to influence its foraging success by attracting prey during both the day and night.     

Weapons or mating signals? Claw shape and mate choice in a fiddler crab

Males often possess traits, such as horns, claws, and tusks, which are used during male–male combat. Studies suggest that selection has fine tuned these weapons to improve their effectiveness and that the shape of these weapons provides cues for males to assess the strength of rivals. Similarly, females might also assess a male’s weaponry to determine his value as a mate. The largest weapon relative to body size is the major claw of male fiddler crabs (genus: Uca). Males use their claws both as a weapon and as a courtship signal, waving it to attract females. We used robotic crabs in two-choice experiments to test female mating preferences based on male claw shape in Uca mjoebergi. First, females did not prefer conspecific claws over any of three alternatively shaped stimuli matched for color and for the rate and pattern of waving. The alternative stimuli were two different heterospecific claws and a plain rectangular shape. Second, females significantly preferred the alternative stimuli when they were presented at a faster wave rate. We conclude that claw shape in U. mjoebergi has not evolved under sexual selection imposed by female mating preferences and is more likely to have evolved under selection for effectiveness during male–male competition.     

Size-based predation by kookaburras (Dacelo novaeguineae) on lizards (Eulamprus tympanum: Scincidae): what determines prey vulnerability?

Lizards and birds are both popular ”model organisms” in behavioural ecology, but the interactions between them have attracted little study. Given the putative importance of birds as predators of diurnal lizards, it is of considerable interest to know which traits (of lizards as well as birds) influence the outcome of a predatory attempt. We studied predation by giant terrestrial kingfishers (kookaburras, Dacelo novaeguineae: Alcedinidae) on heliothermic diurnal lizards (highland water skinks, Eulamprus tympanum: Scincidae), with particular reference to the role of prey (lizard) size. Our approach was twofold: to gather direct evidence (sizes of lizards consumed in the field, compared to those available) and indirect evidence (size-related shifts in lizard behaviour). We quantified the size structure of a natural population of skinks (determined by an extensive mark-recapture program), and compared it to the sizes of wild lizards taken by kookaburras (determined by analysis of prey remains left at the birds’ nests). Kookaburras showed size-based predation: they preyed mainly on small and medium-sized rather than large lizards in the field. However, the mechanism producing this bias remains elusive. It is not due to any distinctive behavioural attributes (locomotor ability, activity level, habitat usage) of the lizards of the size class disproportionately taken by the kookaburras. The greater vulnerability of subadult lizards may reflect subtle ontogenetic shifts in ecological and behavioural traits, but our data suggest that great caution is needed in inferring patterns of vulnerability to predation from indirect measures based on either the prey or the predator alone. Instead, we need direct observations on the interaction between the two. Received: 30 May 2000 / Revised: 29 July 2000 / Accepted: 26 August 2000     

Fracking and indoor radon: Spurious correlation or cause for concern?

Unconventional gas development (fracking) is controversial in large part because of environmental and health concerns. We consider the concern that fracking leads to more carcinogenic radon gas in nearby buildings. Our empirical approach estimates treatment effects where treatment is continuous (number of wells) and varies in intensity (distance to the wells) and in duration of exposure (the time since wells were drilled). The approach allows any potential effect of fracking to vary non-linearly with the distance between the well and test site and, holding distance constant, the time between drilling and testing. Our main model gives a precisely estimated zero effect of wells on radon concentrations in nearby buildings. It also reveals that energy firms drilled wells in places with higher pre-existing radon levels, which, if ignored, makes it appear that wells within 2 km increase indoor radon but wells 3 km away do not. This explains the finding of a prior study showing a link between drilling and indoor radon.     

Metapopulation persistence in a dynamic landscape: more habitat or better stewardship?

Habitat loss and fragmentation has created metapopulations where there were once continuous populations. Ecologists and conservation biologists have become interested in the optimal way to manage and conserve such metapopulations. Several authors have considered the effect of patch disturbance and recovery on metapopulation persistence, but almost all such studies assume that every patch is equally susceptible to disturbance. We investigated the influence of protecting patches from disturbance on metapopulation persistence, and used a stochastic metapopulation model to answer the question: How can we optimally trade off returns from protection of patches vs. creation of patches? We considered the problem of finding, under budgetary constraints, the optimal combination of increasing the number of patches in the metapopulation network vs. increasing the number of protected patches in the network. We discovered that the optimal trade-off is dependent upon all of the properties of the system: the species dynamics, the dynamics of the landscape, and the relative costs of each action. A stochastic model and accompanying methodology are provided allowing a manager to determine the optimal policy for small metapopulations. We also provide two approximations, including a rule of thumb, for determining the optimal policy for larger metapopulations. The method is illustrated with an example inspired by information for the greater bilby, Macrotis lagotis, inhabiting southwestern Queensland, Australia. We found that given realistic costs for each action, protection of patches should be prioritized over patch creation for improving the persistence of the greater bilby during the next 20 years.     

Transatlantic migration by post-breeding puffins: a strategy to exploit a temporarily abundant food resource?

The distribution of Atlantic puffins (Fratercula arctica) from Skellig Michael, south-west Ireland, was investigated using geolocation loggers between the 2010 and 2011 breeding seasons. All tracked birds travelled rapidly west into the North Atlantic at the end of the breeding season in August, with the majority undertaking transatlantic trips from Ireland to the Newfoundland-Labrador shelf. The furthest distance from the colony reached by each bird was not influenced by body mass or sex and was achieved in approximately 20 days. By October, all birds had moved back to the mid Atlantic where they remained resident until returning to the breeding colony. The most parsimonious explanation for the rapid, directed long-distance migration is that birds exploit the seasonally high abundance of prey [e.g., fish species such as capelin (Mallotus villosus) and sandlance (Ammodytes spp.)] off the Canadian coast, which is also utilised by large populations of North American seabirds at this time. Once the availability of this short-term prey resource has diminished, the tracked puffins moved back towards the north-east Atlantic. A relationship between relative abundance of puffins and zooplankton was found in all winter months, but after correcting for spatial autocorrelation, was only significant in November and January. Nevertheless, these results suggest a potential switch in diet from mainly fish during the breeding and early post-breeding periods to zooplankton over the remaining winter period. This study suggests that puffins from south-west Ireland have a long-distance migration strategy that is rare in breeding puffins from the UK and identifies a key non-breeding destination for puffins from Ireland. This has implications for the susceptibility of different breeding populations to the effects of possible climatic or oceanographic change.