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Polymorphic growth in larvae of Maculinea butterflies, as an example of biennialism in myrmecophilous insects.

The presence of annual and biennial individuals within the same population has been recently demonstrated in the myrmecophilous butterflies Maculinea rebeli and Maculinea alcon, which present a cuckoo strategy inside Myrmica nests, and Maculinea arion which is a predatory species. Here, we present field and laboratory data on polymorphic larval growth in two other predatory species of Maculinea: M. teleius and M. nausithous. Body mass distributions of pre-pupation larvae were bimodal in both species. These results point to the existence of larvae that develop in 1 or 2 years. We also showed that the probability of pupation depended on larval body mass. In the case of M. teleius, the critical body mass at which larvae have a 50% probability of pupation is about 80 mg. We suggest that polymorphism in Maculinea may have evolved as an adaptation to life in ant nests, a habitat which protects them from predators and provides food. However, the quality of this resource is highly variable and unpredictable. According to the bet-hedging hypothesis, if the habitat is unpredictable, females should have an advantage by producing more variable offspring. In the case of Maculinea butterflies, this may involve maintaining larvae that develop in 1 or 2 years.

Animals↗

Detecting emergent effects of multiple predator species.

When foraging together, multiple predator species that share a single prey often cause prey mortality that cannot be predicted based on knowledge of predation by each species separately. Modeling and managing the effects of multiple predator species depend on accurately assessing these combined effects. Two methods are currently used to experimentally examine combined predation by multiple predator species: the additive and substitutive experimental designs. I simultaneously employed both experimental designs to examine predation by two crab species on shared mussel prey. I show that the two methods yield results that disagree both quantitatively and qualitatively, leading to very different conclusions about the way that predator species combine to affect prey mortality. This discrepancy occurred because the two methods examine complimentary, but not interchangeable questions. I advocate using an experimental design that incorporates both additive and substitutive designs to achieve a more complete understanding of the combined effects of multiple predator species.

Animals↗

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

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

Animals↗

Environmental variation and the predator-specific responses of tropical stream insects: effects of temperature and predation on survival and development of Australian Chironomidae (Diptera).

The threat posed by predation varies among predator species and with environmental context, and prey species often adjust their responses accordingly. We investigated such effects within an insect assemblage from a tropical Australian stream. These systems are frequently subjected to catastrophic floods, often suggested to reduce the importance of predation in streams, and invertebrate faunas are characterised by relatively broad environmental tolerances. Impacts of the hunting predator Australopelopia prionoptera (Diptera: Chironomidae) and an undescribed ambush predator from the Polycentropodidae (Trichoptera) on survival and development of two species of tubicolous Chironomidae, Echinocladius martini (Orthocladiinae) and Polypedilum australotropicus (Chironominae), were assessed in laboratory microcosms. A further experiment investigated how impacts of Australopelopia varied over a broad range of temperatures, exceeding that experienced annually by the studied populations. Neither predator impacted survivorship for E. martini, but the presence of the polycentropodid caused E. martini to spend longer as larvae and reduced adult longevity, and adult females were smaller-sized and had smaller oocytes. In contrast, both predators reduced survivorship of P. australotropicus, but only Australopelopia affected its development, causing reductions in pupal duration and oocyte size. The observed non-lethal impacts of predation reflect the threat each predator is known to pose to each prey species in situ. Impacts of predation varied little with temperature, reflecting the broad thermal tolerances of all study species. The predator-specific responses of the prey species imply that predation is a significant selective force in tropical Australian streams, although fluctuation in intensity of predation associated with flooding may limit its importance for community structure and prey diversity at larger scales. Our results indicate a more limited scope for environmental modification of predator-prey relationships in faunas characterised by broad physiological tolerances.

Animals↗

Habitat structure, trophic structure and ecosystem function: interactive effects in a bromeliad-insect community.

Although previous studies have shown that ecosystem functions are affected by either trophic structure or habitat structure, there has been little consideration of their combined effects. Such interactions may be particularly important in systems where habitat and trophic structure covary. I use the aquatic insects in bromeliads to examine the combined effects of trophic structure and habitat structure on a key ecosystem function: detrital processing. In Costa Rican bromeliads, trophic structure naturally covaries with both habitat complexity and habitat size, precluding any observational analysis of interactions between factors. I therefore designed mesocosms that allowed each factor to be manipulated separately. Increases in mesocosm complexity reduced predator (damselfly larva) efficiency, resulting in high detritivore abundances, indirectly increasing detrital processing rates. However, increased complexity also directly reduced the per capita foraging efficiency of the detritivores. Over short time periods, these trends effectively cancelled each other out in terms of detrital processing. Over longer time periods, more complex patterns emerged. Increases in mesocosm size also reduced both predator efficiency and detritivore efficiency, leading to no net effect on detrital processing. In many systems, ecosystem functions may be impacted by strong interactions between trophic structure and habitat structure, cautioning against examining either effect in isolation.

Animals↗

Intraguild interactions between spiders and ants and top-down control in a grassland food web.

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis ((15)N and (13)C). Adult wandering spiders were more enriched in (15)N relative to (14)N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed delta(15)N ratios just one trophic level above those of Collembola, and they had similar delta(13)C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.

Animals↗

Population dynamics of thrips prey and their mite predators in a refuge.

Prey refuges are expected to affect population dynamics, but direct experimental tests of this hypothesis are scarce. Larvae of western flower thrips Frankliniella occidentalis use the web produced by spider mites as a refuge from predation by the predatory mite Neoseiulus cucumeris. Thrips incur a cost of using the refuge through reduced food quality within the web due to spider mite herbivory, resulting in a reduction of thrips developmental rate. These individual costs and benefits of refuge use were incorporated in a stage-structured predator-prey model developed for this system. The model predicted higher thrips numbers in presence than in absence of the refuge during the initial phase. A greenhouse experiment was carried out to test this prediction: the dynamics of thrips and their predators was followed on plants damaged by spider mites, either with or without web. Thrips densities in presence of predators were higher on plants with web than on unwebbed plants after 3 weeks. Experimental data fitted model predictions, indicating that individual-level measurements of refuge costs and benefits can be extrapolated to the level of interacting populations. Model-derived calculations of thrips population growth rate enable the estimation of the minimum predator density at which thrips benefit from using the web as a refuge. The model also predicted a minor effect of the refuge on the prey density at equilibrium, indicating that the effect of refuges on population dynamics hinges on the temporal scale considered.

Animals↗

Geographic and within-population variation in the globeflower-globeflower fly interaction: the costs and benefits of rearing pollinators' larvae.

Interspecific interactions can vary within and among populations and geographic locations, and this variation can influence the nature of the interaction (e.g. mutualistic vs. antagonistic) and its evolutionary stability. Globeflowers are exclusively pollinated by flies, whose larvae feed only on their seeds. Here we document geographic variability in costs and benefits in globeflowers in sustaining their pollinating flies throughout the range of this arctic-alpine European plant over several years. A total of 1,710 flower heads from 38 populations were analysed for their carpel, egg and seed contents. Individual and population analyses control for the confounding influences of variation in both: (1) population traits, such as fly density and egg distribution among flower heads; and (2) individuals traits, such as carpel and egg numbers per flower head. Despite considerable variation in ecological conditions and pollinator densities across populations, large proportions (range 33-58%) of seeds were released after predation, with a benefit-to-cost ratio of 3, indicating that the mutualism is stable over the whole globeflower geographical range. The stability of the mutualistic interaction relies on density-dependent competition among larvae co-developing in a flower head. This competition is revealed by a sharp decrease in the number of seeds eaten per larva with increasing larval number, and is intensified by non-uniform egg distribution among globeflowers within a population. Carpel number is highly variable across globeflowers (range 10-69), and flies lay more eggs in large flowers. Most plants within a population contribute to the rearing of pollinators, but the costs are greater for some than for others. Large globeflowers lose more seed to pollinator larvae, but also release more seed than smaller plants. The apparent alignment of interests between fly and plants (positive relationship between numbers of seed released and destroyed) is shown to hide a conflict of interest found when flower size is controlled for.

Analysis of Variance↗

Predator avoidance in phytophagous mites: response to present danger depends on alternative host quality.

We studied whether volatiles released by putative host plants affect the antipredator response of an herbivorous mite, Tetranychus urticae, when the patch was invaded by Phytoseiulus persimilis. Tetranychus urticae laid a lower number of eggs on tomato leaves than on lima bean leaves, suggesting that lima bean is a preferred host food source for T. urticae. In addition, T. urticae preferred lima bean plant volatiles to tomato plant volatiles in a Y-tube olfactometer test. To investigate the antipredator response of T. urticae, we examined the migration of T. urticae from a lima bean leaf disc to a neighbouring leaf disc (either a tomato or lima bean leaf disc) when ten predators were introduced into the original lima bean disc. A Parafilm bridge allowed for migration between the leaf discs. No migrations occurred between leaf discs when there were no predators introduced to the original leaf disc. However, when predators were introduced migrations did occur. When the neighbouring leaf disc was upwind of the original disc, the migration rate of the mite from original lima bean leaf disc to a neighbouring tomato leaf disc was significantly lower than that to a neighbouring lima bean leaf disc. By contrast, when the neighbouring leaf disc was downwind of the original leaf disc, there was no difference in the migration rates between lima bean leaf discs and tomato leaf discs. The number of T. urticae killed by P. persimilis for each treatment was not different, and this clearly shows that the danger was the same in all treatments regardless of the decision made by T. urticae. From these results, we conclude that T. urticae change their antipredator response by evaluating the difference in host plant volatiles in the patch they inhabit.

Analysis of Variance↗

Phenotypically plastic responses of green frog embryos to conflicting predation risk.

Predators have been shown to alter the timing of switch points between life history stages, but few studies have addressed switch point plasticity in prey exposed simultaneously to conflicting predation pressure. We tested hatching responses of green frog (Rana clamitans) embryos subject to perceived predation risk from chemical cues released by two stage-specific predators, predicting that these predators would elicit: (1) directional hatching responses when presented independently, and (2) intermediate phenotypic responses when presented simultaneously. R. clamitans embryos in outdoor exclosures were exposed to cues from an egg predator (freshwater leeches; Nephelopsis obscura), a larval predator (dragonfly nymphs, Aeschna canadensis), and both predators in a 2 x 2 factorial experiment, and changes in hatchling size, hatchling developmental stage, and hatching time were compared to those for control embryos. Leeches alone induced embryos to hatch at a smaller size and an earlier developmental stage than controls, while dragonfly nymphs elicited a delay in egg hatching time that was associated with larger size and later developmental stage at hatching. Embryos failed to respond to simultaneous exposure to both predators, implying that responses to each occurred concurrently and were therefore dampened. Our results indicate that prey under threat from conflicting predators may manifest intermediate defensive phenotypes. Such intermediate responses may result in elevated rates of prey mortality with possible consequences at the population level.

Adaptation, Physiological↗

Behavioural versus physiological mediation of life history under predation risk.

Predator-generated variation in prey energy intake remains the dominant explanation of adaptive response to predation risk in prey life history, morphology and physiology across a wide range of taxa. This "behavioural hypothesis" suggest that chemical or visual signals of predation risk reduce prey energy intake leading to a life history characterized by a small size and late age at maturity. However, size-selective predation can induce either smaller size-early age or large size-late age life history. The alternative "physiological hypothesis" suggests that size-selective cues decouple the relationship between energy and life history, acting instead directly on development. Here we use a series of experiments in a fish-daphnid predator-prey system to ask whether size-selective predator cues induce a physiological mediation of development, overshadowing behaviourally based changes in food intake. We found fish chemical cues reduce the net energy intake in Daphnia magna, suggesting a behaviourally mediated reduction in energy. Experimental manipulation of food levels show further that reductions in food lead to later but smaller size at maturity. However, in line with the physiological hypothesis, we show that D. magna matures earlier and at a smaller size when exposed to fish predation cues. Furthermore, our data shows that they do this by increasing their development rate (earlier maturity) for a given growth rate, resulting in a smaller size at maturity. Our data, from a classic size-selective predation system, indicate that predator-induced changes in this system are driven by physiological mediation of development rather than behavioural mediation of energy intake.

Animals↗

Win-shift and win-stay learning in the short-beaked echidna (Tachyglossus aculeatus).

Numerous previous investigators have explained species differences in spatial memory performance in terms of differences in foraging ecology. In three experiments we attempted to extend these findings by examining the extent to which the spatial memory performance of echidnas (or "spiny anteaters") can be understood in terms of the spatio-temporal distribution of their prey (ants and termites). This is a species and a foraging situation that have not been examined in this way before. Echidnas were better able to learn to avoid a previously rewarding location (to "win-shift") than to learn to return to a previously rewarding location (to "win-stay"), at short retention intervals, but were unable to learn either of these strategies at retention intervals of 90 min. The short retention interval results support the ecological hypothesis, but the long retention interval results do not.

Animals↗

Opportunistic use of cognitive smokescreens by araneophagic jumping spiders.

Little is known about how a prey species' cognitive limitations might shape a predator's prey-capture strategy. A specific hypothesis is investigated: predators take advantage of times when the prey's attention is focussed on its own prey. Portia fimbriata, an araneophagic jumping spider (Salticidae) from Queensland, is shown in a series of 11 experiments to exploit opportunistically a situation in which a web-building spider on which it preys, Zosis genicularis (Uloboridae), is preoccupied with wrapping up its own prey. Experimental evidence supports three conclusions: (1). while relying on optical cues alone, P. fimbriata perceives when Z. genicularis is wrapping up prey; (2). when busy wrapping up prey, the responsiveness of Z. genicularis to cues from potential predators is diminished; and (3). P. fimbriata moves primarily during intervals when Z. genicularis is busy wrapping up prey. P. fimbriata's strategy is effective partly because the wrapping behaviour of Z. genicularis masks the web signals generated by the advancing P. fimbriata's footsteps and also because, while wrapping, Z. genicularis' attention is diverted away from predator-revealing cues.

Animals↗

Interpopulation variation in the risk-related decisions of Portia labiata, an araneophagic jumping spider (Araneae, Salticidae), during predatory sequences with spitting spiders.

The extent to which decision-making processes are constrained in animals with small brains is poorly understood. Arthropods have brains much smaller and simpler than those of birds and mammals. This raises questions concerning limitations on how intricate the decision-making processes might be in arthropods. At Los Baños in the Philippines, Scytodes pallidus is a spitting spider that specialises in preying on jumping spiders, and Portia labiata is a jumping spider that preys on S. pallidus. Scytodid spit comes from the mouth, and egg-carrying females are less dangerous than eggless scytodids because the female uses her chelicerae to hold her eggs. Held eggs block her mouth, and she has to release them before she can spit. The Los Baños P. labiata sometimes adjusts its tactics depending on whether the scytodid encountered is carrying eggs or not. When pursuing eggless scytodids, the Los Baños P. labiata usually takes detour routes that enable it to close in from behind (away from the scytodid's line of fire). However, when pursuing egg-carrying scytodids, the Los Baños P. labiata sometimes takes faster direct routes to reach these safer prey. The Los Baños P. labiata apparently makes risk-related adjustments specific to whether scytodids are carrying eggs, but P. labiata from Sagada in the Philippines (allopatric to Scytodes) fails to make comparable risk-related adjustments.

Animals↗

Bee-eaters ( Merops orientalis) respond to what a predator can see.

Two sets of experiments are reported that show that the small green bee-eater ( Merops orientalis, a small tropical bird) can appreciate what a predator can or cannot see. Bee-eaters avoid entering the nest in the presence of a potential nest predator. In the first set of experiments bee-eaters entered the nest more frequently when the predator was unable to see the nest from its position, as compared to an approximately equidistant position from which the nest could be seen. In the second set of experiments bee-eaters entered the nest more frequently when the predator was looking away from the nest. The angle of gaze from the nest was associated significantly positively with the probability of entering the nest whereas the angle from the bird was not. Birds showed considerable flexibility as well as individual variation in the possible methods of judging the predator's position and direction of gaze.

Animals↗

Profiling transcriptome complexity and secondary metabolite synthesis in a benthic soft coral, Sinularia polydactyla.

Sinularia polydactyla, an abundant Indo-Pacific soft coral species, exhibits biochemical phenotypic plasticity, prompting investigations into differences in mRNA diversity and complexity in response to predation stress. Changes in transcriptome complexity of S. polydactyla cDNA libraries were measured using reannealing rate assays that employ an informatics-based analysis of kinetic profiles. This method allows for quick, high-throughput analysis of sequence complexity and has been used to compare transcriptome-level differences in other marine invertebrates. Here, S. polydactyla colonies were transplanted between two sites exhibiting high and low predation levels. Statistically significant differences between bite scar counts found on different transplant groups suggest site-specific variation in predation. Changes in mRNA pool complexity were quantified to indicate shifts in secondary metabolite concentration between treatment groups. Examining the complexity of the mRNA pool in this soft coral is one of the first steps toward understanding the mechanisms of phenotypic plasticity at a biochemical and molecular level.

Animals↗

Red-tailed monkeys (Cercopithecus ascanius) hunt green pigeons (Treron calva) in the Kalinzu Forest in Uganda.

Red-tailed monkeys (Cercopithecus ascanius) were observed hunting green pigeons (Treron calva) in the Kalinzu Forest in Uganda. During 2 h 39 min, I observed two cases of successful hunting and one case of unsuccessful hunting in a Ficus saussureana tree. Red-tailed monkeys stalked the pigeons until they were within 2-3 m, and then jumped and caught them. In both successful cases, blue monkeys (C. mitis) ran to the hunting site from adjacent trees in order to poach the prey, and the red-tailed monkeys fled. One of these red-tailed monkeys dropped the pigeon while fleeing, and the blue monkey climbed down from the tree to search for it. This is the first record of cercopithecoid monkeys hunting birds that are outside of the nest and moving freely, and also the first record of red-tailed monkeys hunting vertebrates. However rare it may be, the repeated hunting attempts using similar techniques and the immediate attempt of blue monkeys to poach the prey suggest that this type of hunting was not a one-time event that happened by chance. Blue monkeys and an adult chimpanzee (Pan troglodytes schweinfurthii) in and around the fig tree did not attempt to hunt. The hunting of volant birds may be enabled by the small body size and the quick movements of red-tailed monkeys.

Animals↗

Interactions between tayras (Eira barbara) and red-handed howlers (Alouatta belzebul) in eastern Amazonia.

Two tayras (Eira barbara) were observed attacking an infant red-handed howler (Alouatta belzebul) on an island in eastern Brazilian Amazonia, as a nearby adult male watched passively. In a separate incident, four tayras were seen attacking a subadult female on the ground. Tayras were also observed in the vicinity of the two howler study groups on a number of occasions over a 9-month period. During this same period, the two groups lost a third of their members, including infants and subadults, and the remains of six animals were found at the study site. While tayras were the only predators seen attacking the howlers, it remains unclear to what extent they were responsible for apparently high mortality rates in this high-density, isolated howler population.

Alouatta↗