Using stable isotopes to reveal shifts in prey consumption by generalist predators.

The effectiveness of generalist predators in biological control may be diminished if increased availability of alternative prey causes individual predators to decrease their consumption of crop pests. Farming practices that enhance densities of microbidetritivores in the detrital food web can lead to increased densities of generalist predators that feed on pest species. The ability to predict the net biocontrol impact of increased predator densities depends upon knowing the extent to which individual predators may shift to detrital prey and feed less on crop pests when prey of the detritus-based food web are more abundant. We addressed this question by comparing ratios of stable isotopes of carbon (delta13C) and nitrogen (delta15N) in generalist ground predators and two types of prey (crop pests and microbidetritivores) in replicated 8 x 8 m cucurbit gardens subjected to one of two treatments: a detrital subsidy or no addition of detritus (control). Small sheet-web spiders (Linyphiidae) and small wolf spiders (Lycosidae) had delta13C values similar to those of Collembola in both the detrital and control treatments, indicating that small spiders belong primarily to the detrital food web. In control plots the larger generalist predators had delta13C values similar to those of the major insect pests, consistent with their known effectiveness as biocontrol agents. Adding detritus may have caused delta13C of one species of large wolf spider to shift toward that of the microbi-detritivores, although evidence is equivocal. In contrast, another large wolf spider displayed no shift in delta13C in the detrital treatment. Thus, stable isotopes revealed which generalist predators will likely continue to feed on pest species in the presence of greater densities of alternative prey.     

Behavioral response of a generalist predator to chemotactile cues of two taxonomically distinct prey species

Chemotactile cues unintentionally left by animals can play a major role in predator–prey interactions. Specialized predators can use them to find their prey, while prey individuals can assess predation risk. However, little is known to date about the importance of chemotactile cues for generalist predators such as ants. Here, we investigated the response of a generalized predatory ant, Formica polyctena, to cues of two taxonomically distinct prey: a spider (Pisaura mirabilis) and a cricket (Nemobius sylvestris). In analogy, we studied whether crickets and spiders showed antipredator behavior in response to ant cues. When confronted with cues of the two prey species, Formica polyctena workers showed increased residence time and reduced movement speed, which suggests success-motivated searching behavior and thus increased foraging effort. The ants’ response did not differ between cues of the two prey species, coinciding with similar aggression and consumption rates of dead prey. However, the cuticular hydrocarbons, which likely resemble part of the potential cues, differed strongly between the species, with only few methyl-branched alkanes in common. This suggests that ants respond to multiple compounds left by other organisms with prey-search behavior. The two prey species, in turn, showed no detectable antipredator behavior in response to ant cues. Our study shows that ants can detect and respond to chemotactile cues of taxonomically and ecologically distinct prey species, probably to raise their foraging success. Using such chemotactile cues for prey detection may drastically increase their foraging efficiency and thus contribute to the high ecological success of ants.     

Can rare positive interactions become common when large carnivores consume livestock?

Livestock populations in protected areas are viewed negatively because of their interaction with native ungulates through direct competition for food resources. However, livestock and native prey can also interact indirectly through their shared predator. Indirect interactions between two prey species occur when one prey modifies either the functional or numerical responses of a shared predator. This interaction is often manifested as negative effects (apparent competition) on one or both prey species through increased predation risk. But indirect interactions can also yield positive effects on a focal prey if the shared predator modifies its functional response toward increased consumption of an abundant and higher-quality alternative prey. Such a phenomenon between two prey species is underappreciated and overlooked in nature. Positive indirect effects can be expected to occur in livestock-dominated wildlife reserves containing large carnivores. We searched for such positive effects in Acacia-Zizhypus forests of India's Gir sanctuary where livestock (Bubalus bubalis and Bos indicus) and a coexisting native prey (chital deer, Axis axis) are consumed by Asiatic lions (Panthera leo persica). Chital vigilance was higher in areas with low livestock density than in areas with high livestock density. This positive indirect effect occurred because lion predation rates on livestock were twice as great where livestock were abundant than where livestock density was low. Positive indirect interactions mediated by shared predators may be more common than generally thought with rather major consequences for ecological understanding and conservation. We encourage further studies to understand outcomes of indirect interactions on long-term predator-prey dynamics in livestock-dominated protected areas.     

The influence of size-specific indirect interactions in predator-prey systems

Nonlethal indirect interactions between predators often lead to nonadditive effects of predator number on prey survival and growth. Previous studies have focused on systems with at least two different predator species and one prey species. However, most predators undergo extreme ontological changes in phenotype such that interactions between different-sized cohorts of a predator and its prey could lead to nonadditive effects in systems with only two species. This may be important since different-sized individuals of the same species can differ more in their ecology than similar-sized individuals of different species. This study examined trait-mediated indirect effects in a two-species system including a cannibalistic predator with different-sized cohorts and its prey. I tested for these effects using larvae of two stream salamanders, Gyrinophilus porphyriticus (predator) and Eurycea cirrigera (prey), by altering the densities and combinations of predator size classes in experimental streams. Results showed that the presence of large individuals can significantly reduce the impact of density changes of smaller conspecifics on prey survival through nonlethal means. In the absence of large conspecifics, an increase in the relative frequency of small predators significantly increased predation rates, thereby reducing prey survival. However, with large conspecifics present, increasing the density of small predators did not decrease prey survival, resulting in a 14.3% lower prey mortality than predicted from the independent effects of both predator size classes. Small predators changed their microhabitat use in the presence of larger conspecifics. Prey individuals reduced activity in response to large predators but did not respond to small predators. Both predators reduced prey growth. These results demonstrate that the impact of a predator can be significantly altered by two different types of trait-mediated indirect effects in two-species systems: between different-sized cohorts and between different cohorts and prey. This study demonstrates that predictions based on simple numerical changes that assume independent effects of different size classes or ignore size structure can be strongly misleading. We need to account for the size structure within predator populations in order to predict how changes in predator abundance will affect predator-prey dynamics.     

The pit-trapping foraging strategy of the ant lion,Myrmeleon immaculatus DeGeer (Neuroptera: Myrmeleontidae)

Summary Larvae of Myrmeleon immaculatus in large pits captured both large and samll prey, while larvae in small pits captured only the small prey. Larvae in small pits did not respond to large ants, although they always responded by sand-flinging to small ants. Larvae in medium-sized pits often captured large ants only after prolonged and vigorous sand-flipping. Larvae in large pits usually captured large ants with relatively little sand-flipping. Pit enlargement and pit relocation in the laboratory were not significantly correlated with reduction of rations in the first 3 weeks after a pit was built. However, after a month without food, larvae on the average moved once every 10 days, built successively smaller pits, and moved longer distances before building a new pit. In the field pits were dug primarily in response to microclimatological factors and possibly edge-effects. The presence or absence of suitable prey at a site, per se, had no effect on whether or not a larva would dig a pit there. We conclude that these sit-and-wait predators have a relatively large repertoire of behavior that enhances their foraging success, and we contrast it with previously made optimal foraging models relating to pit locations, pit relocations, pit size and ant lion responses.     

Functional responses: a question of alternative prey and predator density

Throughout the study of ecology, there has been a growing realization that indirect effects among species cause complexity in food webs. Understanding and predicting the behavior of ecosystems consequently depends on our ability to identify indirect effects and their mechanisms. The present study experimentally investigates indirect interactions arising between two prey species that share a common predator. In a natural field experiment, we introduced different densities of mealworms (Tenebrio molitor), an alternative prey, to a previously studied predator-prey system in which paper wasps (Polistes dominulus) preyed on shield beetle larvae (Cassida rubiginosa). We tested if alternative prey affects predation on the first prey (i.e., the predator-dependent functional response of paper wasps) by modifying either interference among predators or the effective number of predators foraging on shield beetles. Presence of mealworms significantly reduced the effective number of predators, whereas predator interference was not affected. In this way, the experimentally introduced alternative prey altered the wasps' functional response and thereby indirectly influenced C. rubiginosa density. In all prey-density combinations offered, paper wasps constantly preferred T. molitor. This led to an asymmetrical, indirect interaction between both prey species: an increase in mealworm density significantly relaxed predation on C. rubiginosa, whereas an increase in C. rubiginosa density intensified predation on mealworms. Such asymmetrical outcomes of a fixed food preference can significantly affect the population dynamics of the species involved. In spite of the repeated finding of a Type III functional response in this system, our experiment did not reveal switching behavior in paper wasps. The variety of mechanisms underlying direct and indirect interactions within our study system exemplifies the importance of incorporating alternative prey when investigating the impact of a generalist predator on a focal prey population under realistic field conditions.     

Critical patch sizes for food-web modules

Because patch size and connectivity may strongly impact the assemblage of species that occur on a patch, the types of food-web interactions that occur among those species may also depend on spatial structure. Here, we identify whether food-web interactions among salt-marsh-inhabiting arthropods vary with patch size and connectivity, and how such changes in trophic structure might feed back to influence the spatial distribution of prey. In a multiyear survey, patch-restricted predators exhibited steeper occupancy-patch-size relationships than herbivores, and species' critical patch sizes were correlated with overall rarity. As a result, the presence of food-web modules depended strongly on patch size: large and well-connected patches supported complex food-web modules, but only the simplest modules involving the most abundant species were found on small patches. Habitat-generalist spiders dominated on small patches, and predation pressure from such species may contribute to the observed lower densities of mesopredators on small patches. Overall, patch size and connectivity influenced the types of modules present on a patch through differential loss of rare, patch-restricted predators, but predation by generalist predators may be a key mechanism influencing the spatial structure of certain prey species.     

Lake to land subsidies: experimental addition of aquatic insects increases terrestrial arthropod densities

Aquatic insects are a common and important subsidy to terrestrial systems, yet little is known about how these inputs affect terrestrial food webs, especially around lakes. Myvatn, a lake in northern Iceland, has extraordinary midge (Chironomidae) emergences that result in large inputs of biomass and nutrients to terrestrial arthropod communities. We simulated this lake-to-land resource pulse by collecting midges from Myvatn and spreading their dried carcasses on 1-m2 plots at a nearby site that receives very little midge deposition. We hypothesized a positive bottom-up response of detritivores that would be transmitted to their predators and would persist into the following year. We sampled the arthropod community once per month for two consecutive summers. Midge addition resulted in significantly different arthropod communities and increased densities of some taxa in both years. Detritivores, specifically Diptera larvae, Collembola, and Acari increased in midge-addition plots, and so did some predators and parasitoids. Arthropod densities were still elevated a year after midge addition, and two years of midge addition further increased the density of higher-order consumers (e.g., Coleoptera and Hymenoptera). Midge addition increased arthropod biomass by 68% after one year and 108% after two years. By manipulating the nutrient pulse delivered by midges we were able to elucidate food web consequences of midge deposition and spatial and temporal dynamics that are difficult to determine based on comparative approaches alone. Resources cross ecosystem boundaries and are assimilated over time because of life-history strategies that connect aquatic and terrestrial food webs and these systems cannot be fully understood in isolation from each other.     

The disruption of an ant-aphid mutualism increases the effects of birds on pine herbivores

Predators affect herbivores directly and indirectly, by consumptive and nonconsumptive effects, and the combined influence of multiple predators is shaped by interactions among predators. I documented the individual and combined effects of birds (chickadees, nuthatches, warblers) and ants (Formica podzolica) on arthropods residing in pine (Pinus ponderosa) canopies in a factorial field experiment. Birds and ants removed herbivores but simultaneously benefited them by removing predatory arthropods. Birds and ants had net negative and positive effects, respectively, on the abundance of herbivore prey, supporting the notion that vertebrate predators have stronger negative effects on herbivores than do arthropod predators. Aphids (ant-tended and untended species) constituted three-quarters of herbivore biomass. The effect of birds on ant-tended aphids was twice that on untended aphid species or tended aphid species without ants. This was not due to there being more ant-tended aphids for birds to prey on; tended and untended aphid species were in similar abundances in the absence of birds. Instead, the effects of birds were strengthened by attributes of the mutualism that rendered tended aphids susceptible to predation. These dynamics led to nonadditive effects of birds and ants: birds only reduced tended aphid species and total herbivore abundances on trees with ants, while ants only increased tended aphid species and total herbivore abundances in the absence of birds. Consequently, top predators in this system only influenced total herbivore abundance when they disrupted an ant-aphid mutualism.     

Social spiders catch larger prey: a study of Anelosimus eximius (Araneae: Theridiidae)

Summary During a 1-year-study in tropical Panama, prey of the social theridiid Anelosimus eximius was analysed at two locations and compared with the potential prey spectrum according to sweepnet catches, pitfall traps and bowl traps. Compared with other web-building spiders, A. eximius catch an unusually high number of large insects: about 90% are flying ants, beetles,lepidopterans hemipterans, cockroaches and grasshoppers. This is the result of a communal strategy to overwhelm prey. Webs are maintained commonly, and several spiders attack an entangled insect simultaneously. More spiders participate on insects that are larger and struggle more. The ability to catch large prey insects is discussed as a major driving factor for sociality in spiders.     

Indirect effects of prey swamping: differential seed predation during a bamboo masting event

Resource pulses often involve extraordinary increases in prey availability that "swamp" consumers and reverberate through indirect interactions affecting other community members. We developed a model that predicts predator-mediated indirect effects induced by an epidemic prey on co-occurring prey types differing in relative profitability/preference and validated our model by examining current-season and delayed effects of a bamboo mass seeding event on seed survival of canopy tree species in mixed Patagonian forests. The model shows that predator foraging behavior, prey profitability, and the scale of prey swamping influence the character and strength of short-term indirect effects on various alternative prey. When in large prey-swamped patches, nonselective predators decrease predation on all prey types. Selective predators, instead, only benefit prey of similar quality to the swamping species, while very low or high preference prey remain unaffected. Negative indirect effects (apparent competition) may override such positive effects (apparent mutualism), especially for highly preferred prey, when prey-swamped patches are small enough to allow predator aggregation and/or predators show a reproductive numerical response to elevated food supply. Seed predation patterns during bamboo (Chusquea culeou) masting were consistent with predicted short-term indirect effects mediated by a selective predator foraging in large prey-swamped patches. Bamboo seeds and similarly-sized Austrocedrus chilensis (ciprés) and Nothofagus obliqua (roble) seeds suffered lower predation in bamboo flowered than nonflowered patches. Predation rates on the small-seeded Nothofagus dombeyi (coihue) and the large-seeded Nothofagus alpina (rauli) were independent of bamboo flowering. Indirect positive effects were transient; three months after bamboo seeding, granivores preyed heavily upon all seed types, irrespective of patch flowering condition. Moreover, one year after bamboo seeding, predation rates on the most preferred seed (rauli) was higher in flowered than in nonflowered patches. Despite rapid predator numerical responses, short-term positive effects can still influence community recruitment dynamics because surviving seeds may find refuge beneath the litter produced by bamboo dieback. Together, our theoretical analysis and experiments indicate that indirect effects experienced by alternative prey during and after prey-swamping episodes need not be universal but can change across a prey quality spectrum, and they critically depend on predator-foraging rules and the spatial scale of swamping.     

Long-term aboveground and belowground consequences of red wood ant exclusion in boreal forest

Despite their ubiquity, the role of ants in driving ecosystem processes both aboveground and belowground has been seldom explored, except within the nest. During 1995 we established 16 ant exclusion plots of approximately 1.1 x 1.1 m, together with paired control plots, in the understory layer of a boreal forest ecosystem in northern Sweden that supports high densities of the mound-forming ant Formica aquilonia, a red wood ant species of the Formica rufa group. Aboveground and belowground measurements were then made on destructively sampled subplots in 2001 and 2008, i.e., 6 and 13 years after set-up. While ant exclusion had no effect on total understory plant biomass, it did greatly increase the relative contribution of herbaceous species, most likely through preventing ants from removing their seeds. This in turn led to higher quality resources entering the belowground subsystem, which in turn stimulated soil microbial biomass and activity and the rates of loss of mass and carbon (C) and nitrogen (N) from litter in litterbags placed in the plots. This was accompanied by losses of approximately 15% of N and C stored in the humus on a per area basis. Ant exclusion also had some effects on foliar stable isotope ratios for both C and N, most probably as a consequence of greater soil fertility. Further, exclusion of ants had multitrophic effects on a microbe-nematode soil food web with three consumer trophic levels and after six years promoted the bacterial-based relative to the fungal-based energy channel in this food web. Our results point to a major role of red wood ants in determining forest floor vegetation and thereby exerting wide-ranging effects on belowground properties and processes. Given that the boreal forest occupies 11% of the Earth's terrestrial surface and stores more C than any other forest biome, our results suggest that this role of ants could potentially be of widespread significance for biogeochemical nutrient cycling, soil nutrient capital, and sequestration of belowground carbon.     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Indirect effects of an exploited predator on recruitment of coral-reef fishes

The more ecologists examine the role of trait-mediated indirect interactions (TMIIs), especially in regulating predator-prey interactions, the more we recognize their fundamental role in structuring food webs. However, most empirical evidence for TMIIs comes from studies that are either conducted in laboratory or mesocosm venues or are restricted to simple food webs involving lower trophic-level animals. Here, I quantified the direct and indirect effects of interactions between high-level vertebrate predators on their vertebrate prey using a field experiment. Specifically, I tested how varying densities of a large-bodied, top predator (Nassau grouper; Epinephelus striatus) affected persistence, growth, and behavior of two smaller-bodied, intermediate predators (coney and graysby groupers; Cephalopholis fulva and C. cruentata) on 20 isolated patch reefs in the Bahamas. Large-bodied groupers are capable of consuming their smaller-bodied counterparts, and previous observational studies have indicated that local abundances of these groupers are negatively correlated. I measured the effects of interactions among groupers on lower trophic-level prey by quantifying recruitment of coral-reef fishes to the reefs. The field experiment demonstrated a strong trophic cascade that was entirely mediated by modified behavior of the intermediate predators. These results indicate that indirect, nonlethal interactions in natural systems can have strong cascading effects even at high trophic levels and in high-diversity food webs. Incorporating the complexity of such indirect effects into fisheries management may improve the sustainability of fished populations and strengthen marine conservation efforts; however these results also indicate that the effects of fishing are complex and difficult to predict.     

Bottom-up effects of plant genotype on aphids, ants, and predators

Theory predicts that bottom-up ecological forces can affect community dynamics, but whether this extends to the effects of heritable plant variation on tritrophic communities is poorly understood. In a field experiment, I contrasted the effects of plant genotype (28 genotypes; 1064 plants), aphid density, and the presence/absence of mutualistic ants in affecting the per capita population growth of a specialist aphid herbivore, as well as the effects of plant genotype on the third trophic level. Plant genotype strongly affected aphid population growth rate, explaining 29% of the total variation in growth rate, whereas aphid density and ant-aphid interactions explained substantially less variation (< 2%) in aphid population growth rate. Plant genotype also had direct and indirect effects on the third trophic level, affecting the abundance of aphid-tending ants and the richness of predators. Multiple regression identified several heritable plant traits that explained 49% of the variation in aphid growth rate and 30% of the variation in ant abundance among plant genotypes. These bottom-up effects of plant genotype on tritrophic interactions were independent of the effects of either initial aphid density or the presence/absence of mutualistic ants. This study shows that plant genotype can be one of the most important ecological factors shaping tritrophic communities.     

Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

Field evidence for pervasive indirect effects of fishing on prey foraging behavior

The indirect, ecosystem-level consequences of ocean fishing, and particularly the mechanisms driving them, are poorly understood. Most studies focus on density-mediated trophic cascades, where removal of predators alternately causes increases and decreases in abundances of lower trophic levels. However, cascades could also be driven by where and when prey forage rather than solely by prey abundance. Over a large gradient of fishing intensity in the central Pacific's remote northern Line Islands, including a nearly pristine, baseline coral reef system, we found that changes in predation risk elicit strong behavioral responses in foraging patterns across multiple prey fish species. These responses were observed as a function of both short-term ("acute") risk and longer-term ("chronic") risk, as well as when prey were exposed to model predators to isolate the effect of perceived predation risk from other potentially confounding factors. Compared to numerical prey responses, antipredator behavioral responses such as these can potentially have far greater net impacts (by occurring over entire assemblages) and operate over shorter temporal scales (with potentially instantaneous response times) in transmitting top-down effects. A rich body of literature exists on both the direct effects of human removal of predators from ecosystems and predators' effects on prey behavior. Our results draw together these lines of research and provide the first empirical evidence that large-scale human removal of predators from a natural ecosystem indirectly alters prey behavior. These behavioral changes may, in turn, drive previously unsuspected alterations in reef food webs.     

Reciprocal subsidies between freshwater and terrestrial ecosystems structure consumer resource dynamics

Cross-ecosystem movements of material and energy, particularly reciprocal resource fluxes across the freshwater-land interface, have received major attention. Freshwater ecosystems may receive higher amounts of subsidies (i.e., resources produced outside the focal ecosystem) than terrestrial ecosystems, potentially leading to increased secondary production in freshwaters. Here we used a meta-analytic approach to quantify the magnitude and direction of subsidy inputs across the freshwater-land interface and to determine subsequent responses in recipient animals. Terrestrial and freshwater ecosystems differed in the magnitude of subsidies they received, with aquatic ecosystems generally receiving higher subsidies than terrestrial ecosystems. Surprisingly, and despite the large discrepancy in magnitude, the contribution of these subsidies to animal carbon inferred from stable isotope composition did not differ between freshwater and terrestrial ecosystems, likely due to the differences in subsidy quality. The contribution of allochthonous subsidies was highest to primary consumers and predators, suggesting that bottom-up and top-down effects may be affected considerably by the input of allochthonous resources. Future work on subsidies will profit from a food web dynamic approach including indirect trophic interactions and propagating effects.     

Balancing between mutualism and exploitation: the symbiotic interaction between Lasius ants and aphids

Nests of Lasius niger (L.) ants were given varied food regimens to test whether their behaviour towards an aphid partner, Aphis fabae (Scop.), changed with alternative food supplies. Honeydew collection and predation on aphids were measured by video monitoring the movement of ants between their nest and an aphid aggregation. Data collected from the aphid aggregations enabled comparisons between remaining aphid biomass and between the tending intensities of the ants. I tested how ant behaviour was influenced by their access to alternative prey and sugar. The results showed that ants accepted a honey solution as a substitute for the honeydew produced by aphids. Ants not only attended their aphid partners, but also preyed on them. The average predation rate increased eightfold when ants were offered the alternative of sugar, whereas alternative prey had no significant effect. In contrast, ant-tending intensity decreased with alternative sugar whereas alternative prey elicited no effect.     

Disentangling trophic relationships in a High Arctic tundra ecosystem through food web modeling

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator-prey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.