Effects of behavioral and morphological plasticity on risk of predation in a Neotropical tadpole

Predator-induced phenotypic plasticity is widespread among aquatic animals, however the relative contributions of behavioral and morphological shifts to reducing risk of predation remain uncertain. We tested the phenotypic plasticity of a Neotropical tadpole (Rana palmipes) in response to chemical cues from predatory Belostoma water bugs, and how phenotype affects risk of predation. Behavior, morphology, and pigmentation all were plastic, resulting in a predator-induced phenotype with lower activity, deeper tail fin and muscle, and darker pigmentation. Tadpoles in the predator cue treatment also grew more rapidly, possibly as a result of the nutrient subsidy from feeding the caged predator. For comparison to phenotypes induced in the experiment, we quantified the phenotype of tadpoles from a natural pool. Wild-caught tadpoles did not match either experimentally induced phenotype; their morphology was more similar to that produced in the control treatment, but their low swimming activity was similar to that induced by predator cues. Exposure of tadpoles from both experimental treatments and the natural pool to a free-ranging predator confirmed that predator-induced phenotypic plasticity reduces risk of predation. Risk of predation was comparable among wild-caught and predator-induced tadpoles, indicating that behavioral shifts can substantially alleviate risk in tadpoles that lack the typical suite of predator-induced morphological traits. The morphology observed in wild-caught tadpoles is associated with rapid growth and high competition in other tadpole species, suggesting that tadpoles may profitably combine a morphology suited to competition for food with behaviors that minimize risk of predation.     

Fitness and community consequences of avoiding multiple predators

We investigated the fitness and community consequences of behavioural interactions with multiple predators in a four-trophic-level system. We conducted an experiment in oval flow-through artificial-stream tanks to examine the single and interactive sublethal effects of brook trout and stoneflies on the size at emergence of Baetis bicaudatus (Ephemeroptera: Baetidae), and the cascading trophic effects on algal biomass, the food resource of the mayflies. No predation was allowed in the experiment, so that all effects were mediated through predator modifications of prey behaviour. We reared trout stream Baetis larvae from just before egg development until emergence in tanks with four treatments: (1) water from a holding tank with two brook trout (trout odour), (2) no trout odour + eight stoneflies with glued mouthparts, (3) trout odour + stoneflies and (4) no trout odour or stoneflies. We ended the experiment after 3 weeks when ten male and ten female subimagos had emerged from each tank, measured the size of ten male and ten female mature nymphs (with black wing pads), and collected algal samples from rocks at six locations in each tank. To determine the mechanism responsible for sublethal and cascading effects on lower trophic levels we made day and night observations of mayfly behaviour for the first 6 days by counting mayflies drifting in the water column and visible on natural substrata in the artificial streams. Trout odour and stoneflies similarly reduced the size of male and female Baetis emerging from artificial streams, with non-additive effects of both predators. While smaller females are less fecund, a fitness cost of small male size has not been determined. The mechanism causing sublethal effects on Baetis differed between predators. While trout stream Baetis retained their nocturnal periodicity in all treatments, stoneflies increased drift dispersal of mayflies at night, and trout suppressed night-time feeding and drift of mayflies. Stoneflies had less effect on Baetis behaviour when fish odour was present. Thus, we attribute the non-additivity of effects of fish and stoneflies on mayfly growth to an interaction modification whereby trout odour reduced the impact of stoneflies on Baetis behaviour. Since stonefly activity was also reduced in the presence of fish odour, this modification may be attributed to the effect of fish odour on stonefly behaviour. Only stoneflies delayed Baetis emergence, suggesting that stoneflies had a greater sublethal effect on Baetis fitness than did trout. Delayed emergence may reduce Baetis fitness by increasing risks of predation and parasitism on larvae, and increasing competition for mates or oviposition sites among adults. Finally, algal biomass was higher in tanks with both predators than in the other three treatments. These data implicate a behavioural trophic cascade because predators were not allowed to consume prey. Therefore, differences in algal biomass were attributed to predator-induced changes in mayfly behaviour. Our study demonstrates the importance of considering multiple predators when measuring direct sublethal effects of predators on prey fitness and indirect effects on lower trophic levels. Identification of an interaction modification illustrates the value of obtaining detailed information on behavioural mechanisms as an aid to understanding the complex interactions occurring among components of ecological communities. Received: 20 March 1997 / Accepted: 29 September 1997     

The dynamics of predation risk assessment: responses of anuran larvae to chemical cues of predators

1. While the antipredator behaviour of prey has been well studied, little is known about the rules governing the predation risk assessment of prey. In this study, I measured the activity levels of predator-naive green frog (Rana clamitans) tadpoles during and after exposures to the chemical cue of predatory larval dragonflies (Anax spp.). I then used the lengths of the time lags from the end of the cue exposures until the tadpoles returned to a control level of activity as an index of the perceived risk of the tadpoles. 2. While tadpoles always responded upon exposure to the Anax chemical cue by strongly reducing their activity level, their perceived risk increased asymptotically over time during the initial period of the cue exposure. Tadpoles of all size classes perceived increasing risk in proportion to chemical cue concentration, but the length of time that tadpoles responded during cue exposure and the length of their post-exposure time lags decreased with increasing body mass. 3. The results suggest that the perceived risk of green frog tadpoles varies over time and does not correspond directly to their behavioural response (i.e. activity level). However, their perceived risk does appear to vary in accordance with the predation risk associated with the Anax chemical cue and the reliability of the information from the cue, and therefore may be predictable.     

Foraging time and spatial patterns of predation in experimental populations

Summary Responses of the predaceous mites Phytoseiulus persimilis, Typhlodromus (=Metaseiulus) occidentalis, and Amblyseius andersoni to spatial variation in egg density of the phytophagous mite, Tetranychus urticae, were studied in the laboratory.The oligophagous predator P. persimilis showed initially a direct density dependent foraging time allocation and variation in foraging time increased with prey density. With changes in prey density due to predation, predator foraging rates (per hour) decreased with time and density dependent foraging gradually became density independence, because P. persimilis continued to respond to initial prey density, instead of the changing prey density and distribution. The consequent spatial pattern of predation by P. persimilis was density independent, although slopes of predation rate-prey density regressions increased with time.Compared with P. persimilis, the narrowly polyphagous predator T. occidentalis responded relatively slowly to the the presence or absence of prey eggs but not to prey density: the mean and variation of foraging time spent in patches with prey did not differ with prey density, but was significantly greater in patches with prey eggs than in patches without eggs. Prey density and distribution changed only slightly due to predation and overall foraging rates remained more or less constant. The consequent spatial pattern of predation by T. occidentalis was inversely density dependent. As with P. persimilis, slopes of predation rate-prey density regressions increased with time (i.e. the inverse density dependence in T. occidentalis became weaker through time).The broadly polyphagous predator A. andersoni showed density independent foraging time allocation with variation independent of prey density. With changes in prey density over time due to prey depletion, overall foraging rates decreased. The consequent spatial pattern of predation by A. andersoni also changed through time; it initially was inversely density dependent, but soon became density independent.Overall, P. persimilis and T. occidentalis spent more time in prey patches than A. andersoni, suggesting that A. andersoni tended to spend more time moving outside patches. The overall predation rates and searching efficiency were higher in P. persimilis than in A. andersoni and T. occidentalis. Predator reproduction was highest in P. persimilis, lower in T. occidentalis and the lowest A. andersoni.The differences in response to prey distribution among the three predaceous species probably reflect the evolution of these species in environments with different patterns of prey distribution. The degree of polyphagy is a major determinant of the aggregative response, but other attributes such as handling time are also important in other aspects of phytoseiid foraging behavior (e.g. searching efficiency or predation rate).     

Direct and indirect effects of predators on the dominant invertebrates of two freshwater littoral communities

Summary Two congeneric damselfly species, Enallagma traviatum and E. aspersum, dominate the littoral macroinvertebrates of Bays Mountain Lake and of the adjacent fish-free Ecology Pond, respectively (northeastern Tennessee, USA). Extending previous experimental studies, we test seven hypotheses concerning the role of fish (bluegill sunfish, Lepomis macrochirus) and larvaldragonfly (Anax junius) predation, competitive effects on damselflies, and the interaction between competition and predation, in determining invertebrate dominance in these communities. Three types of experiments were conducted: an enclosure experiment within Ecology Pond, an outdoor replicated tub experiment, and a laboratory behavior experiment. The in-situ enclosure experiment showed that E. traviatum larvae were more susceptible to Anax predation than were E. aspersum larvae; a tendency toward greater vulnerability to fish of E. aspersum compared with E. traviatum was not statistically significant. The outdoor tub experiment confirmed both of these trends with statistically significant results. In the tubs, both predators inhibited feeding of both zygopterans (as indicated by reduced fecal mass), particularly for E. aspersum in the presence of fish. This effect appears to have been primarily indirect, mediated through exploitation of the zooplankton. We also detected competitive effects of E. traviatum on E. aspersum: E. traviatum reduced the emergence and increased the exposure above the substrate of E. aspersum. In the absence of predators, E. traviatum inhibited feeding of E. aspersum via interference. In the laboratory behavior experiment, predators inhibited crawling by E. aspersum. E. aspersum was more exposed than was E. traviatum; it swam and crawled more than did E. traviatum, considerably increasing these movements at night. Over all, E. traviatum consistently appeared to be the more cryptic of the two species, and E. aspersum appeared to be much more active. Our results suggest an explanation for the clear difference in structure between communities like Bays Mountain Lake and Ecology Pond: predaceous fish eliminate large invertebrate predators and shift the community toward cryptic forms at relatively low densities, reflecting the effects of both predation and exploitation competition. In the absence of fish, large invertebrate predators are less able to deplete littoral invertebrates but may favor the more active forms, perhaps because these are better able to avoid invertebrate predators.     

The effect of larval predators Thanasimus dubius (Coleoptera: Cleridae), produced by an improved system of rearing, against the southern pine beetle Dendroctonus frontalis (Coleoptera: Curculionidae)

The southern pine beetle, Dendroctonus frontalis (Coleoptera: Curculionidae) (SPB), is known to be a major bark beetle pest of pines throughout the southeastern United States. A common predator of bark beetles, Thanasimus dubius (Coleoptera: Cleridae), has been suggested to play a prevalent role on SPB dynamics. Evaluations of T. dubius have been limited by rearing methods; an artificial diet for larval T. dubius exists, and preservatives such as sorbic acid could help to maximize diet shelf-life and enhance the efficiency of the rearing system. The effects of sorbic acid at different concentrations (0%, 0.1% and 0.2%) in the larval diet for T. dubius were measured, and the effects of increased feeding time intervals (2-3 vs. 5 days) on predator performance evaluated. In addition, an experimental bioassay was conducted where newly hatched T. dubius larvae were released at four densities (0, 50, 100, and 200 per log) on pine logs infested by SPB. Sorbic acid in the diet reduced female fecundity (by 20-40%), but did not affect adult T. dubius size or longevity. However, using this preservative may not be necessary because it had no effect on the overall efficiency of the rearing system, while refreshing the larval diet every 5 days (compared with 2-3 days) did improve its efficiency, even without sorbic acid. The release of larval T. dubius resulted in a highly significant effect on the SPB ratio of increase (RI). This experiment was facilitated by the improvements in our rearing methods.     

The arthropod predators of ant-mimetic and aposematic prey: a serological analysis

Abstract.

• 1 ELISA (enzyme-linked immunosorbant assay) was used to identify the arthropod predators of two common herbivores of a desert lupine: the aposematic plant bug Lopidea nigridea and the ant-mimetic plant bug Coquillettia insignis.
• 2 Despite the fact that the prey are closely related, the two antisera were sufficiently specific to distinguish between the antigen against which they were formed and the antigen of the other species: both antisera were 3 times as reactive against their homologous antigen as they were against their heterologous antigen.
• 3 In tests on gut contents of field-collected predators, ELISA results were generally consistent with laboratory no-choice data for the five most common arthropod species on lupine: there was a strong correlation between attack rates for each predator-prey pair as measured in the laboratory, and per cent positive reactions in ELISA as measured using field-collected predators. In particular, L.nigridea antiserum was shown to be significantly less reactive against the crab spider Xysticus montanensis when compared to C.insignis antiserum, paralleling results of laboratory no-choice experiments.
• 4 The usefulness of serological analysis as a tool for arthropod trophic link identification is discussed.

     

Mechanics of the inner ear of the bullfrog (Rana catesbeiana): the contact membranes and the periotic canal

The frog inner ear consists of a complex of fluid-filled membranous sacs and canals containing eight distinct clusters of sensory hair cells. In this study we attempt to delineate the potential pathways for acoustic energy flow toward two of these clusters located within the amphibian papilla and the basilar papilla. Detailed morphological measurements of the periotic canal based on internal casts of the inner ear in the bullfrog (Rana catesbeiana) revealed that it is divided into a wide, tapered section and a narrower section comprised of two branches – one short and blind projecting into the endolymphatic space and another longer, terminating in the round window. Additionally, we used laser Doppler velocimetry to record the velocity responses of the contact membranes of the amphibian papilla and basilar papilla. We found that the acoustic energy flow through these two structures is frequency dependent such that the amphibian papilla contact membrane displays a peak velocity amplitude at frequencies less than 500 Hz, whereas the basilar papilla contact membrane velocity response exhibits a maximum above 1100 Hz. Our data advocate a mechanical substrate underlying the frequency segregation in the auditory nerve fibers innervating the amphibian papilla and the basilar papilla. Accepted: 9 March 2000     

Predator-induced morphological changes in an amphibian: predation by dragonflies affects tadpole shape and color

Predator-induced defenses are well studied in plants and invertebrate animals, but have only recently been recognized in vertebrates. Gray treefrog (Hylachrysoscelis) tadpoles reared with predatory dragonfly (Aeshnaumbrosa) larvae differ in shape and color from tadpoles reared in the absence of dragonflies. By exposing tadpoles to tail damage and the non-lethal presence of starved and fed dragonflies, we determined that these phenotypic differences are induced by non-contact cues present when dragonflies prey on Hyla. The induced changes in shape are in the direction that tends to increase swimming speed; thus, the induced morphology may help tadpoles evade predators. Altering morphology in response to predators is likely to influence interactions with other species in the community as well. Received: 17 April 1996 / Accepted: 18 September 1996     

Interspecific variation in the escape responses of aphids: effect on risk of predation from foliar-foraging and ground-foraging predators

A series of laboratory experiments was conducted to determine the effect of interspecific differences on prey defensive behavior on the susceptibility of two aphid species (Acyrthosiphon pisum and A. kondoi) to a ground-foraging predator, Harpalus pennsylvanicus, and a foliar-foraging predator, Coccinella septempunctata. These organisms are representative of a biologically and economically important predator/prey system in alfalfa. The primary defensive behavior of both aphid species toward C. septempunctata was to “drop” from the plant. Both aphid species were significantly more likely to drop from the plant in the presence of C. septempunctata. However, when C. septempunctata was present, a significantly lower proportion of A. kondoi individuals dropped (0.42 ± 0.07) compared to A. pisum (0.73 ± 0.08). As a result of their lower propensity to drop from the plant A. kondoi individuals are significantly more likely to be consumed by C. septempunctata. Conversely, the higher propensity of A. pisum individuals to drop increased their susceptibility to ground-foraging predators. When A. pisum was the prey species, ground-foraging predators made a significant contribution to overall aphid suppression and there was a significant synergistic interaction between ground and foliar-foraging predators. When A. kondoi was the prey there was no interaction between the predator species. As either a cause or consequence of its higher propensity to drop, A. pisum seems to be more adapted for survival and dispersal off the plant. In comparison to A. kondoi individuals, A. pisum individuals relocate plants more quickly (63 ± 41 s vs. 164 ± 39 s), disperse farther (18 ± 1.7 cm vs. 13 ± 0.66 cm), and survive longer (37 ± 2.0 h vs. 25 ± 2.0) off the plant. This study demonstrates the importance of prey defensive behavior in determining the susceptibility of a prey species to a multiple-predator complex. Received: 24 February 1997 / Accepted:17 December 1997     

Additive interactions between the moon snail Euspira heros and the sea star Asterias forbesi, two predators of the surfclam Spisula solidissima

A laboratory experiment was conducted to determine whether the sea star Asterias forbesi and the naticid gastropod Euspira heros feed on surfclams, Spisula solidissima, in an additive or non-additive manner. Predators were allowed to feed on clams with conspecifics and in the presence of the other predator species. Clam mortality (measured as the rate of decline of clam number) and predator feeding rates were noted. To determine the effects of temperature on interactions among the predators, the experiment was conducted at three different temperatures. At all temperatures, feeding rate of each predator was not affected by the presence of the other species, and clam mortality in the presence of both predators was predictable from mortality in the presence of a single predator species. These additive interactions are most likely a result of habitat partitioning between the predators, with naticid snails being infaunal and sea stars being epifaunal. Previous studies in a variety of systems show no clear pattern of occurrence of non-additive interactions. Relatively small differences in predator or prey behavior may be responsible for the presence or absence of non-additive interactions. Received: 6 August 1998 / Accepted: 25 January 1999     

Native species vulnerability to introduced predators: testing an inducible defense and a refuge from predation

To manage the impacts of biological invasions, it is important to determine the mechanisms responsible for the effects invasive species have on native populations. When predation by an invader is the mechanism causing declines in a native population, protecting the native species will involve elucidating the factors that affect native vulnerability. To examine those factors, this study measured how a native species responded to an introduced predator, and whether the native response could result in a refuge from predation. Predation by the green crab, Carcinus maenas, has contributed to the decline in numbers of native soft-shell clams, Mya arenaria, and efforts to eradicate crabs have proven futile. We tested how crab foraging affected clam burrowing, and how depth in the sediment affected clam survival. Clams responded to crab foraging by burrowing deeper in the sediment. Clams at shallow depths were more vulnerable to predation by crabs. Results suggest soft-shell clam burrowing is an inducible defense in response to green crab predation because burrowing deeper results in a potential refuge from predation by crabs. For restoring the native clam populations, tents could exclude crabs and protect clams, but when tents must be removed, exposing the clams to cues from foraging crabs should induce the clams to burrow deeper and decrease vulnerability. In general, by exposing potential native prey to cues from introduced predators, we can test how the natives respond, identify whether the response results in a potential refuge, and evaluate the risks to native species survival in invaded communities.     

Optimal foraging when predation risk increases with patch resources: an analysis of pollinators and ambush predators

Pollinators and their predators share innate and learned preferences for high quality flowers. Consequently, pollinators are more likely to encounter predators when visiting the most rewarding flowers. I present a model of how different pollinator species can maximize lifetime resource gains depending on the density and distribution of predators, as well as their vulnerability to capture by predators. For pollinator species that are difficult for predators to capture, the optimal strategy is to visit the most rewarding flowers as long as predator density is low. At higher predator densities and for pollinators that are more vulnerable to predator capture, the lifetime resource gain from the most rewarding flowers declines and the optimal strategy depends on the predator distribution. In some cases, a wide range of floral rewards provides near‐maximum lifetime resource gains, which may favor generalization if searching for flowers is costly. In other cases, a low flower reward level provides the maximum lifetime resource gain and so pollinators should specialize on less rewarding flowers. Thus, the model suggests that predators can have qualitatively different top‐down effects on plant reproductive success depending on the pollinator species, the density of predators, and the distribution of predators across flower reward levels.     

Identification of carbonic anhydrase activity in bullfrog olfactory receptor neurons: histochemical localization and role in CO2 chemoreception

The objectives of this study were to determine: (1) the frequency and distribution of carbonic anhydrase (CA) activity in the bullfrog nasal cavities, and (2)␣whether inhibition of nasal CA affects the olfactory receptor response to CO₂ or other odorants. It was found, using Hansson's staining technique, that some olfactory receptor neurons exhibited CA activity and that these CA-positive receptors were distributed throughout the nasal cavity with peak densities in the dorsal and ventral sensory epithelial regions. To test for the role of CA in olfactory transduction, electro-olfactograms (EOGs) were recorded from the surface of the ventral sensory epithelium in response to 2-s pulses of 5% CO₂ and amyl acetate before and after topical CA inhibition with acetazolamide (10⁻³mol · l⁻¹). In 52 bullfrogs, 1222 sites on the ventral epithelium were tested resulting in 23 locations that exhibited a response to 5% CO₂. Inhibition of CA caused an immediate 65% reduction in the EOG response to CO₂ while the response to amyl acetate was not affected. These results, along with the histochemical localization of CA in some olfactory receptor neurons, indicate that CA plays a role in the detection of CO₂ in frog olfactory neurons and that only a small population of olfactory receptor neurons are CO₂ sensitive. Accepted: 31 July 1997     

Direct and indirect effects of predation on mosquitofish behavior and survival

Predation can have strong direct and indirect effects on the behavior of prey. We investigated whether predation by chain pickerel (Esox niger) caused adult eastern mosquitofish (Gambusia holbrooki) to alter their habitat use and whether pickerel predation influenced survival of adult and neonate mosquitofish. The number of adult mosquitofish using the riskier of three habitats was lowest when two predators occupied the risky habitat, intermediate in the treatment with one predator, and highest when no predators occurred there. More mosquitofish neonates survived high predation treatments than treatments lacking pickerel. We conclude that pickerel predation causes adult mosquitofish to shift to refuge habitats. The pattern of neonate survival suggests that adult habitat use may create a refuge from cannibalism for neonate mosquitofish, resulting in higher neonate survival in treatments with more pickerel. Hence, pickerel predation has a direct effect on adult mosquitofish behavior and a strong indirect effect on neonate survival. Both interspecific and intraspecific predation can effect prey populations and can interact to produce important indirect effects.     

Control and interaction of the cardiovascular and respiratory systems in anuran amphibians

In anuran amphibians, respiratory rhythm is generated within the central nervous system (CNS) and is modulated by chemo- and mechanoreceptors located in the vascular system and within the CNS. The site for central respiratory rhythmogenesis and the role of various neurotransmitters and neuromodulators is described. Ventilatory air flow is generated by a positive pressure, buccal force pump driven by efferent motor output from cranial nerves. The vagus (cranial nerve X) also controls heart rate and pulmocutaneous arterial resistance that, in turn, affect cardiac shunts within the undivided anuran ventricle; however, little is known about the control of central vagal motor outflow to the heart and pulmocutaneous artery. Anatomical evidence indicates a close proximity of the centers responsible for respiratory rhythmogenesis and the vagal motoneurons involved in cardiovascular regulation. Furthermore, anurans in which phasic feedback from chemo- and mechanoreceptors is prevented by artificial ventilation exhibit cardiorespiratory interactions that appear similar to those of conscious animals. These observations indicate interactions between respiratory and cardiovascular centers within the CNS. Thus, like mammals and other air-breathing vertebrates, the cardio-respiratory interactions in anurans result from both feedback and feed-forward mechanisms.     

Type and nature of cues used by Nucella lapillus to evaluate predation risk

The ability of prey to detect and adequately respond to predation risk influences immediate survival and overall fitness. Chemical cues are commonly used by prey to evaluate risk, and the purpose of this study was to elicit the nature of cues used by prey hunted by generalist predators. Nucella lapillus are common, predatory, intertidal snails that evaluate predatory risk using chemical cues. Using Nucella and a suite of its potential predators as a model system, we explored how (1) predator type, (2) predator diet, and (3) injured conspecifics and heterospecifics influence Nucella behavior. Using laboratory flumes, we determined that Nucella responded only to the invasive green crab (Carcinus maenas), the predator it most frequently encounters. Nucella did not respond to rock crabs (Cancer irroratus) or Jonah crabs (Cancer borealis), which are sympatric predators but do not frequently encounter Nucella because these crabs are primarily subtidal. Predator diet did not affect Nucella responses to risk, although starved predator response was not significantly different from controls. Since green crabs are generalist predators, diet cues do not reflect predation risk, and thus altering behavior as a function of predator diet would not likely benefit Nucella. Nucella did, however, react to injured conspecifics, a strategy that may allow them to recognize threats when predators are difficult to detect. Nucella did not react to injured heterospecifics including mussels (Mytilus edulis) and herbivorous snails Littorina littorea, suggesting that they are responding to chemical cues unique to their species. The nature of cues used by Nucella allows them to minimize costs associated with predator avoidance.     

Predictable changes in predation mortality as a consequence of changes in food availability and predation risk

Theory predicts that animals will have lower activity levels when either the risk of predation is high or the availability of resources in the environment is high. If encounter rates with predators are proportional to activity level, then we might expect predation mortality to be affected by resource availability and predator density independent of the number of effective predators. In a factorial experiment, we tested whether predation mortality of larval wood frogs, Rana sylvatica, caused by a single larval dragonfly, Anax junius, was affected by the presence of additional caged predators and elevated resource levels. Observations were consistent with predictions. The survival rate of the tadpoles increased when additional caged predators were present and when additional resources were provided. There was no significant interaction term between predator density and food concentration. Lower predation rates at higher predator density is a form of interference competition. Reduced activity of prey at higher predator density is a potential general mechanism for this widespread phenomenon. Higher predation rates at low food levels provides an indirect mechanism for density-dependent predation. When resources are depressed by elevated consumer densities, then the higher activity levels associated with low resource levels can lead to a positive association between consumer density and consumer mortality due to predation. These linkages between variation in behaviour and density-dependent processes argue that variation in behaviour may contribute to the dynamics of the populations. Because the capture rate of predators depends on the resources available to prey, the results also argue that models of food-web dynamics will have to incorporate adaptive variation in behaviour to make accurate predictions.     

The influence of salt and nitrogen on herbivore abundance: direct and indirect effects

We report on the influence of experimentally increased interstitial salinity and plant nitrogen on the abundance of the delphacid planthopper, Prokelisia marginata (Van Duzee) (Homoptera: Delphacidae), which feeds on salt marsh cordgrass, Spartina alterniflora. We also report the effects of these treatments on parasitism of P. marginata eggs by the fairyfly parasitoid, Anagrus sophiae (Hymenoptera: Mymaridae). Soil salinity was significantly elevated following the addition of salt pellets broadcast over the ground and plant foliar nitrogen was significantly increased after the addition of fertilizer. The addition of fertilizer increased P. marginata densities on Spartina but addition of salt did not. Neither treatment significantly affected levels of egg parasitism by A. sophiae. In this system direct effects of plants on their herbivores via changes in plant chemistry appear more important than indirect effects of plants on herbivores via their natural enemies. Received: 1 August 1997 / Accepted: 29 September 1997     

Nematode and Grape Rootstock Interactions Including an Improved Understanding of Tolerance

Sixteen cultivars of grape were screened over a two-year period in the presence or absence of 10 different nematode populations. Populations of Meloidogyne spp., Xiphinema index, and Mesocriconema xenoplax developed more rapidly and caused greater damage than populations of X. americanum and Tylenchulus semipenetrans. Populations of mixed Meloidogyne spp. having a history of feeding on grape were among the fastest developing populations. Tolerance to nematode parasitism appeared to be based on different mechanisms. Slow developing, less pathogenic nematode populations often stimulated vine growth, thus vines appeared to possess tolerance. Likewise, cultivars selected for nematode resistance often stimulated vine growth when fed upon by the nematode. However, tolerance sources that resulted from nematode resistance are vulnerable due to the occurrence of populations that break resistance mechanisms. Growth of cultivars with phylloxera (Daktalospharia vitifoliae) resistance was unchanged by the presence of nematodes, indicating that phylloxera resistance may provide a useful source of nematode relief. These and several additional sources of specific tolerance are discussed.