Construction and evaluation of a robust trophic network model for the northern Gulf of Mexico ecosystem

A critical issue in understanding trophic connectivity in ecological systems is the lack in quality and quantity of information about feeding habits. In this work, we present a method for integrating a diversity of feeding habits data from published studies to evaluate the impact on indices that describe characteristics of individual taxa and their connectivity. We focus our study on feeding habits of the fishes of the northern Gulf of Mexico and seek to understand the importance of the forage fish Gulf Menhaden (Brevoortia patronus) in predator diets. We created a database of diet studies from the northern Gulf of Mexico that included six diet metrics: frequency of occurrence, wet weight, dry weight, number, volume, index of relative importance, and index of caloric importance. We then used this information to construct a set of traditional networks (all prey and predators were from a single taxonomic level and trophic connections were parameterized with a single diet metric). We also constructed a “robust” network where all taxa were identified to the lowest taxonomic level and trophic connections were parameterized using a resampling approach that included all available information. Linear regression and resampling methods were used to convert data reported in other diet metrics into the frequency of occurrence diet metric. For both traditional and robust networks, we used network indices to describe topological properties. With the robust network, we conducted removal simulations where the forage fish species Gulf Menhaden, and associated Clupeidae representatives, were removed from the network and the feeding effort of the predators was reallocated among their other prey items. We found that network and node-specific indices were sensitive to the choice of taxonomy and diet metric level. In the robust network, predator species with the greatest number of identified prey had the lowest precision in their connections and prey from the Arthropoda phyla had the lowest precision for connections. From the removal and reallocation simulations, we found that Actinopterygii and Arthropoda were the most impacted prey taxa with 1.2% to 4.3% increase in predation and approximately 23 taxa would receive 50% of the reallocated predation. Overall, the resampling methods we present provide a potential means for combining disparate diet data and enables a comprehensive understanding of trophic interactions within an ecosystem.     

THE TROPHIC ROLE OF MARINE MAMMALS IN THE NORTHERN GULF OF ST. LAWRENCE

The trophic role of apex predators was evaluated in the northern Gulf of St. Lawrence ecosystem. An Ecopath model was developed for the period 1985–1987 prior to the collapse of commercially exploited demersal fish stocks in this area. Marine mammal trophic levels were estimated by the model at 4.1 for cetaceans, 4.4 for harp seals (Pagophilus groenlandicus), 4.7 for hooded seals (Cystophora cristata), 4.5 for gray seals (Halichoerus grypus), and 4.3 for harbor seals (Phoca vitulina). Harp seals were the third most important predator on vertebrate prey following large Atlantic cod (Gadus morhua) and redfish (Sebastes spp.). Different seal species preyed on different levels of the food chain. Harp seals preyed on most trophic groups, whereas larger seals, such as gray seals and hooded seals, mainly consumed higher trophic levels. The model suggested that apex predators had a negative effect on their dominant prey, the higher trophic level fish, but an indirect positive feedback on the prey of their preferred prey, mainly American plaice (Hippoglossoides platessoides), flounders, skates, and benthic invertebrates. Our results suggest that both marine mammals and fisheries had an impact on the trophic structure.     

Induction of phenotypic plasticity in rattlesnake trophic morphology by diet manipulation

Gape‐limited predators are restricted in the shape and size of prey items they can ingest by their trophic morphology. Evolutionary theory predicts that gape‐limited predators, such as rattlesnakes, should possess plasticity in their trophic morphology to allow them to respond to environmental cues about their prey base. This study examined the effects of two possible influences over trophic morphology in the pit‐viper Crotalus viridis viridis. Snakes from six litters were exposed to diet manipulations performed over 480 days. By day 480, snakes from two prey‐size treatments exhibited significantly different head shapes. Snakes reared on whole rodents had broader heads, whereas snakes force‐fed homogenized prey had narrower heads. Shape differences varied among litters, suggesting that not all litters responded the same to diet manipulations. Results suggest that trophic morphology of rattlesnakes is plastic, at least in some litters, and can be induced by prey items. J. Morphol. 275:1339–1348, 2014. © 2014 Wiley Periodicals, Inc.     

Do juvenile gape-limited predators compensate for their small size when feeding?

When juvenile and adult animals occur syntopically, juveniles are at a distinct performance disadvantage due to their absolutely small size. Yet, optimal foraging theory predicts that juvenile predators should feed efficiently in order to compete with adults for food, and to minimize their exposure to predators. Previous authors have suggested that one way for juvenile animals to accomplish these ecological tasks is by increasing their overall feeding performance relative to adults (compensation hypothesis). Nonetheless, only a handful of studies have tested whether juvenile animals have increased feeding performance (e.g. decreased ingestion and/or handling times relative to body size) compared with adults. We tested this hypothesis by examining the ontogeny of head dimensions and feeding performance (ingestion time and number of mandibular protractions) on fish prey for broad-banded water snakes Nerodia fasciata . Individuals were fed fish scaled in a 1:1 ratio to their head width. All head dimensions scaled with significant negative allometry versus body size, and thus smaller snakes had relatively larger heads for their body size compared with larger snakes. By contrast, most head variables (except head volume) exhibited positive allometry versus head length, demonstrating that larger snakes had larger head dimensions relative to head size compared with smaller snakes. In the performance trials, smaller snakes had worse feeding performances when feeding on similarly sized fish prey (relative to their head width) compared with larger snakes. Therefore, these data show that smaller water snakes do not compensate for their size through increased feeding performance.     

Critical indirect effects of climate change on sub‐Antarctic ecosystem functioning

Sub‐Antarctic islands represent critical breeding habitats for land‐based top predators that dominate Southern Ocean food webs. Reproduction and molting incur high energetic demands that are sustained at the sub‐Antarctic Prince Edward Islands (PEIs) by both inshore (phytoplankton blooms; “island mass effect”; autochthonous) and offshore (allochthonous) productivity. As the relative contributions of these sustenance pathways are, in turn, affected by oceanographic conditions around the PEIs, we address the consequences of climatically driven changes in the physical environment on this island ecosystem. We show that there has been a measurable long‐term shift in the carbon isotope signatures of the benthos inhabiting the shallow shelf region of the PEIs, most likely reflecting a long‐term decline in enhanced phytoplankton productivity at the islands in response to a climate‐driven shift in the position of the sub‐Antarctic Front. Our results indicate that regional climate change has affected the balance between allochthonous and autochthonous productivity at the PEIs. Over the last three decades, inshore‐feeding top predators at the islands have shown a marked decrease in their population sizes. Conversely, population sizes of offshore‐feeding predators that forage over great distances from the islands have remained stable or increased, with one exception. Population decline of predators that rely heavily on organisms inhabiting the inshore region strongly suggest changes in prey availability, which are likely driven by factors such as fisheries impacts on some prey populations and shifts in competitive interactions among predators. In addition to these local factors, our analysis indicates that changes in prey availability may also result indirectly through regional climate change effects on the islands' marine ecosystem. Most importantly, our results indicate that a fundamental shift in the balance between allochthonous and autochthonous trophic pathways within this island ecosystem may be detected throughout the food web, demonstrating that the most powerful effects of climate change on marine systems may be indirect.     

Relative strengths of trait-mediated and density-mediated indirect effects of a predator vary with resource levels in a freshwater food chain

Predators can affect the density and traits (e.g. morphology, behavior) of their prey, and either change may influence how prey interact with their resources. Thus, predators can interact indirectly with resource species (i.e. two trophic levels below) through two separate mechanisms. The relative strengths of these two kinds of indirect effects have rarely been compared directly, and how their relative importance varies across environmental gradients is virtually unknown. We investigated the relative strength of trait- and density-mediated indirect effects of the predatory insect Belostoma flumineum on algal communities through predation on the pond snail, Physa gyrina , across a gradient of basal resource abundance. Because prey balance the benefits of foraging against the increased risk of predation while foraging, the availability of the prey's resource should influence the strength of anti-predator behavioral responses and hence the strength of trait-mediated indirect interactions. Belostoma presence had positive indirect effects on resources as expected and total predator effects were constant across the basal resource gradient. At low initial resource levels, trait-mediated indirect effects on algal biomass exceeded density-mediated indirect effects, while at high initial resources the reverse was true. Snails showed similar habitat use across the resource gradient suggesting that the anti-predator response was most likely a depression of activity levels.     

Trophic cascades in rocky shore tide pools: distinguishing lethal and nonlethal effects

The effects of predators on the density of their prey can have positive indirect effects on the abundance of the preys resource via a trophic cascade. This concept has strongly influenced contemporary views of how communities are structured. However, predators also can transmit indirect effects by inducing changes in prey traits. We show that the mere presence of predator risk cues can initiate a trophic cascade in rocky shore tide pools. In large (mean surface area =9 m²), natural tide pools, we manipulated crab density and their foraging ability to examine the relative importance of lethal (density-mediated) and non-lethal (trait-mediated) predator effects to algal community development. We found that perceived predation risk reduced snail density as much as the direct predation treatment, showing that green crab predation was not an important factor regulating local snail density. Instead, snail emigration away from resident crabs appears to be the most important factor regulating local snail density. As a result, the abundance of ephemeral green algae was similar in the predation risk and direct predation treatments, suggesting that the consumption of snails by crabs plays a minimal role in mediating the trophic cascade. Increased attention to trait-mediated effects that are transmitted by predator-induced changes in prey behavior may change our view of how predators exert their strong influence on community structure.     

Ontogenetic shift in diet of a large elapid snake is facilitated by allometric change in skull morphology

Evolutionary Ecology - As snakes are limbless, gape-limited predators, their skull is the main feeding structure involved in prey handling, manipulation and feeding. Ontogenetic changes in prey...     

Predators and Resources Influence Phosphorus Transfer along an Invertebrate Food Web through Changes in Prey Behaviour

Predators play a fundamental role in prey trophic behaviour, with indirect consequences for species coexistence and ecosystem functioning. Resource quality and availability also influence prey trophic behaviour, with potential effects on predator-prey dynamics. Although many studies have addressed these topics, little attention has been paid to the combined effects of predators and resources on prey species coexistence and nutrient transfer along food chains, especially in detritus-based systems. To determine the influence of predators and resource quality on the movement and P uptake of detritivores, we carried out a field experiment on the River Kelvin (Scotland) using ³²P to test the hypothesis of reduced prey vagility among resource patches as a strategy to avoid predation. Thirty leaf sacks containing alder leaves and two detritivore prey populations (Asellus aquaticus and Lymnaea peregra) were placed in cages, half of them with two predator species (Dendrocoelum lacteum and Erpobdella octoculata) and the other half without predators. Five alder leaf bags, each individually inoculated with a different fungus strain to simulate a patchy habitat, were placed inside each leaf sack. One bag in each sack was labelled with ³²P, in order to assess the proportion of detritivores using it as food and thus their movement among the five resource patches. Three replicates for each labelled fungus and each predation treatment (i.e. with and without predators) were left on the riverbed for 7 days. The presence of predators had negligible effects on the number of detritivores in the leaf bags, but it did reduce the proportion of ³²P-labelled detritivores and their P uptake. The most strongly affected species was A. aquaticus, whose vagility, trophic overlap with L. peregra and P uptake were all reduced. The results confirm the importance of bottom-up and top-down forces acting simultaneously to regulate nutrient transfer along food chains in patchy habitats.     

Incorporating anthropogenic effects into trophic ecology: predator–prey interactions in a human-dominated landscape

Apex predators perform important functions that regulate ecosystems worldwide. However, little is known about how ecosystem regulation by predators is influenced by human activities. In particular, how important are top-down effects of predators relative to direct and indirect human-mediated bottom-up and top-down processes? Combining data on species'' occurrence from camera traps and hunting records, we aimed to quantify the relative effects of top-down and bottom-up processes in shaping predator and prey distributions in a human-dominated landscape in Transylvania, Romania. By global standards this system is diverse, including apex predators (brown bear and wolf), mesopredators (red fox) and large herbivores (roe and red deer). Humans and free-ranging dogs represent additional predators in the system. Using structural equation modelling, we found that apex predators suppress lower trophic levels, especially herbivores. However, direct and indirect top-down effects of humans affected the ecosystem more strongly, influencing species at all trophic levels. Our study highlights the need to explicitly embed humans and their influences within trophic cascade theory. This will greatly expand our understanding of species interactions in human-modified landscapes, which compose the majority of the Earth''s terrestrial surface.     

Wait Until Dark? Daily Activity Patterns and Nest Predation by Snakes

Predation involves costs and benefits, so predators should employ tactics that reduce their risk of injury or death and that increase their success at capturing prey. One potential way that predators could decrease risk and increase benefits is by attacking prey at night when risks may be reduced and prey more vulnerable. Because some snakes are facultatively nocturnal and prey on bird nests during the day and night, they are ideal for assessing the costs and benefits of diurnal vs. nocturnal predation. We used automated radiotelemetry and cameras to investigate predation on nesting birds by two species of snakes, one diurnal and the other facultatively nocturnal. We predicted that snakes preying on nests at night should experience less parental nest defence and capture more adults and nestlings. Rat snakes (Pantherophis obsoletus) were relatively inactive at night (23–36% activity) but nearly always preyed on nests after dark (80% of nest predations). Conversely, racers (Coluber constrictor) were exclusively diurnal and preyed on nests during the times of day they were most active. These results are consistent with rat snakes strategically using their capacity for facultative nocturnal activity to prey on nests at night. The likely benefit is reduced nest defence because birds defended their nests less vigourously at night. Consistent with nocturnal predation being safer, rat snake predation events lasted three times longer at night than during the day (26 vs. 8 min). Nocturnal nest predation did not make nests more profitable by increasing the likelihood of capturing adults or removing premature fledging of nestlings. The disconnect between rat snake activity and timing of nest predation seems most consistent with rat snakes locating prey during the day using visual cues but waiting until dark to prey on nests when predation is safer, although designing a direct test of this hypothesis will be challenging.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Convergence in trophic morphology and feeding performance among piscivorous natricine snakes

Piscivory has independently evolved numerous times amongst snakes, and therefore these animals provide a powerful opportunity to test for convergent evolution in a vertebrate feeding system. In this study, we integrate performance trials with comparative methods to test the hypothesis that piscivory drives convergence in trophic morphology and feeding performance among natricine snakes. Within and across species, increasing the relative length of the quadrate bone in the skull is positively and strongly linked to a reduction in the time needed to swallow large fish prey. Thus, our feeding experiments suggest that a longer quadrate bone enables snakes to better conform their head shape to the shape of the prey during swallowing. Ancestral diet reconstructions and phylogenetically corrected multiple regression analyses further reveal that evolutionary increases in piscivory are coupled to the evolution of relatively longer quadrates, and hence improved feeding performance on fish prey in these animals. The exploitation of similar dietary niches drives the evolution of convergent trophic morphologies and feeding performances in natricine snakes.     

Income breeding allows an aquatic snake Seminatrix pygaea to reproduce normally following prolonged drought-induced aestivation

1. Capital breeding is an ideal reproductive strategy for many ectotherms because it provides a disassociation between feeding and reproduction, a necessary requirement for animals that become anorexic during pregnancy. Among ectotherms, some viviparous snakes (e.g. Viperidae) exemplify the capital breeding strategy because many species (i) do not feed during pregnancy due to behavioural conflicts between reproduction and foraging, and (ii) take more than one season to accumulate sufficient energetic stores for reproduction. 2. Isolated wetlands often exhibit extreme annual fluctuations in environmental conditions with prolonged droughts periodically leaving wetlands completely dry and devoid of prey. Following droughts, however, wetlands can be extremely productive, rendering prey resources virtually unlimited for some species. 3. This study examines drought survival strategy and reproductive ecology of a small aquatic snake Seminatrix pygaea (Cope) in an isolated wetland. Seminatrix pygaea are atypical from most sympatric snake species in that (i) their small body size, reliance on aquatic prey, and high rates of evaporative water loss make them ill-suited to overland movement, and (ii) they may not be subject to costs typically associated with feeding during pregnancy. 4. We hypothesized that S. pygaea would survive periodic multiyear droughts by aestivating within the dried wetland, a survival strategy heretofore undocumented in snakes. Further, we hypothesized that if S. pygaea rely on 'typical' snake reproductive strategies of 'adaptive anorexia' and capital breeding, reproductive output would be reduced in the first wet year following drought. 5. By encircling a 10-ha wetland with a continuous drift fence before it refilled we were able to demonstrate that S. pygaea were present within the dried wetland prior to the onset of spring rains that refilled the wetland in 2003. Our results suggest that S. pygaea are capable of surviving multiyear droughts by aestivating within the dried wetland. 6. Despite having presumably depleted energy reserves during the drought, S. pygaea reproduced with the same frequency and fecundity during the first season following refilling of the wetland as in pre-drought years. 7. The ability of S. pygaea to rebound rapidly from the stresses of prolonged drought is due in part to their reproductive ecology. Seminatrix pygaea readily feed throughout pregnancy and consequently can rapidly translate high prey abundances into reproductive output through income breeding.     

Predator‐driven elemental cycling: the impact of predation and risk effects on ecosystem stoichiometry

Empirical evidence is beginning to show that predators can be important drivers of elemental cycling within ecosystems by propagating indirect effects that determine the distribution of elements among trophic levels as well as determine the chemical content of organic matter that becomes decomposed by microbes. These indirect effects can be propagated by predator consumptive effects on prey, nonconsumptive (risk) effects, or a combination of both. Currently, there is insufficient theory to predict how such predator effects should propagate throughout ecosystems. We present here a theoretical framework for exploring predator effects on ecosystem elemental cycling to encourage further empirical quantification. We use a classic ecosystem trophic compartment model as a basis for our analyses but infuse principles from ecological stoichiometry into the analyses of elemental cycling. Using a combined analytical‐numerical approach, we compare how predators affect cycling through consumptive effects in which they control the flux of nutrients up trophic chains; through risk effects in which they change the homeostatic elemental balance of herbivore prey which accordingly changes the element ratio herbivores select from plants; and through a combination of both effects. Our analysis reveals that predators can have quantitatively important effects on elemental cycling, relative to a model formalism that excludes predator effects. Furthermore, the feedbacks due to predator nonconsumptive effects often have the quantitatively strongest impact on whole ecosystem elemental stocks, production and efficiency rates, and recycling fluxes by changing the stoichiometric balance of all trophic levels. Our modeling framework predictably shows how bottom‐up control by microbes and top‐down control by predators on ecosystems become interdependent when top predator effects permeate ecosystems.     

Variation in the diet of the viperine snake Natvix mama in relation to prey availability

The diet of the viperine snake was compared with food availability in the Ebro Delta, a wetland largely occupied by rice fields, in 1990 and 1991. Snake selection of prey type and size was studied seasonally and by snake group: males, females and immature snakes. Overall, feeding activity (percentage of individuals with prey and number of prey per stomach) increased with food availability. Diet analysis showed that viperine snakes mainly foraged on the green frog Rana perezi (adults and tadpoles) and the carp Cyprinus earpio. Conversely, viperine snakes rejected the mosquito fish Gambusia holbroki which is the most abundant species in autumn, when Natrix maura has a low feeding activity. Statistical comparisons between viperine snake diet and prey availability showed that males selected small carp, immature snakes selected tadpoles and, in spring, females selected frogs. The selection of small carp by males may reflect a sexual divergence of trophic niche related to sexual size dimorphism, as females are larger than males. As tadpoles are presumably easier to catch than fish, tadpole selection by immature individuals may reflect variance in capture abilities. In spring, the selection of frogs by females overlapped with vitellogenesis, suggesting that females compensate for the cost of reproduction by selecting green frogs, which have a greater biomass and higher energy content than fish. Carps eaten in spring were smaller than in summer. Moreover, in summer viperine snakes selected smaller carp than the available mean size. This divergent tendency between carp size selection and carp size availability reveals how seasonal diet shifts in prey size selection may be a response to an increase in prey size.     

Animal-Borne Imaging Reveals Novel Insights into the Foraging Behaviors and Diel Activity of a Large-Bodied Apex Predator,the American Alligator (Alligator mississippiensis)

Large-bodied, top- and apex predators (e.g., crocodilians, sharks, wolves, killer whales) can exert strong top-down effects within ecological communities through their interactions with prey. Due to inherent difficulties while studying the behavior of these often dangerous predatory species, relatively little is known regarding their feeding behaviors and activity patterns, information that is essential to understanding their role in regulating food web dynamics and ecological processes. Here we use animal-borne imaging systems (Crittercam) to study the foraging behavior and activity patterns of a cryptic, large-bodied predator, the American alligator (Alligator mississippiensis) in two estuaries of coastal Florida, USA. Using retrieved video data we examine the variation in foraging behaviors and activity patterns due to abiotic factors. We found the frequency of prey-attacks (mean = 0.49 prey attacks/hour) as well as the probability of prey-capture success (mean = 0.52 per attack) were significantly affected by time of day. Alligators attempted to capture prey most frequently during the night. Probability of prey-capture success per attack was highest during morning hours and sequentially lower during day, night, and sunset, respectively. Position in the water column also significantly affected prey-capture success, as individuals’ experienced two-fold greater success when attacking prey while submerged. These estimates are the first for wild adult American alligators and one of the few examples for any crocodilian species worldwide. More broadly, these results reveal that our understandings of crocodilian foraging behaviors are biased due to previous studies containing limited observations of cryptic and nocturnal foraging interactions. Our results can be used to inform greater understanding regarding the top-down effects of American alligators in estuarine food webs. Additionally, our results highlight the importance and power of using animal-borne imaging when studying the behavior of elusive large-bodied, apex predators, as it provides critical insights into their trophic and behavioral interactions.     

Prey size diversity hinders biomass trophic transfer and predator size diversity promotes it in planktonic communities

Body size exerts multiple effects on plankton food-web interactions. However, the influence of size structure on trophic transfer remains poorly quantified in the field. Here, we examine how the size diversity of prey (nano-microplankton) and predators (mesozooplankton) influence trophic transfer efficiency (using biomass ratio as a proxy) in natural marine ecosystems. Our results support previous studies on single trophic levels: transfer efficiency decreases with increasing prey size diversity and is enhanced with greater predator size diversity. We further show that communities with low nano-microplankton size diversity and high mesozooplankton size diversity tend to occur in warmer environments with low nutrient concentrations, thus promoting trophic transfer to higher trophic levels in those conditions. Moreover, we reveal an interactive effect of predator and prey size diversities: the positive effect of predator size diversity becomes influential when prey size diversity is high. Mechanistically, the negative effect of prey size diversity on trophic transfer may be explained by unicellular size-based metabolic constraints as well as trade-offs between growth and predation avoidance with size, whereas increasing predator size diversity may enhance diet niche partitioning and thus promote trophic transfer. These findings provide insights into size-based theories of ecosystem functioning, with implications for ecosystem predictive models.     

Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Relative strengths of top-down and bottom-up forces in a tropical forest community

We tested integrative bottom-up and top-down trophic cascade hypotheses with manipulative experiments in a tropical wet forest, using the ant-plant Piper cenocladum and its associated arthropod community. We examined enhanced nutrients and light along with predator and herbivore exclusions as sources of variation in the relative biomass of plants, their herbivores (via rates of herbivory), and resident predaceous ants. The combined manipulations of secondary consumers, primary consumers, and plant resources allowed us to examine some of the direct and indirect effects on each trophic level and to determine the relative contributions of bottom-up and top-down cascades to the structure of the community. We found that enhanced plant resources (nutrients and light) had direct positive effects on plant biomass. However, we found no evidence of indirect (cascading through the herbivores) effects of plant biomass on predators or top predators. In contrast, ants had indirect effects on plant biomass by decreasing herbivory on the plants. This top-down cascade occurred whether or not plant resources were enriched, conditions which are expected to modify top-down forces. Received: 9 August 1998 / Accepted: 1 December 1998